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JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

VoIume 23

Supplement 3

ON THE FREQUENCY OF BUTTERFLIES IN EASTERN BRAZIL, WITH A LIST OF THE BUTTERFLY FAUNA OF POCOS de CALDAS, MINAS GERAIS

Heinz Ebert

Faculdade de Filosofia, Ciencias e Letras de Rio Claro S. Paulo, Brazil1

With the aid of the "Fundacao de Amparo a" Pesquisa do Estado de Sao Paulo" and of the "Conselh6 Nacional de Pesquisas."

Table of Contents

Page

I.    Introduction---------------------------------------------   1

II.   Types of butterfly frequency-----------------------------   1

III.  The regional (extensive) frequency-----------------------   2

IV.   The individual (intensive) frequency---------------------   4

V.    The daily frequency of species---------------------------   6

VI.   The seasonal variation of the daily frequency of species--   8

VII.  The daily frequency of the total number of individuals---   14

VIII. The seasonal variation of the total number of individuals-   15

IX.   Conclusions----------------------------------------------   20

X.    Additional observations

(a)  Influence of the number of observers-----------------   21

(b)  Influence of amount of collecting hours--------------   22

(c) Observations on annual frequency of species numbers—   27

(d) Observations in the high Tropics---------------------   28

Literature cited-----------------------------------------------   30

Appendix I. Extensive frequency of the different families of

butterflies in three selected areas of Eastern

Brazil---------------------------------------------   31

Appendix II. List of the 572 species of butterflies found at

Pocos de Caldas (Minas Gerais) during 37 collecting

days----------------------------------------------   35

Frequency of Butterflies in Eastern Brazil 1

I. INTRODUCTION

Since C. L. Remington (1955) first recorded his highest catch of butterfly species in one day, from the eastern part of the U.S.A., various contributions have been published on the same subject in The Journal of the Lepidoptevists r Society: Hemming (1955) and Langer (1955) from the French Alps, Wiltshire (1956) from Persia, Shull (1958, 1962) from northern India. As is to be expected, the highest number from any Hoiarctic locality (50 to 60) is lower than the value obtained in the Palaeotropics (101), although comparison is difficult because the last value was the result of a party of seven collectors, whereas all others refer to one observer only. Nothing from the Neotropics has been published on this subject.

The present writer paid special attention to problems of butterfly frequency from the fall of 1950 through 1966. After every field trip any species met with and identified was noted in a card file, with its individual frequency, using five ranks from very rare to abundant. Also, since July, 1958, the number of species, grouped into subfamilies, was noted in a special table. The present data (through December, 1966) result from 212 full-day excursions, nine from the Amazonian region and 203 from eastern Brazil. Only the latter are taken into consideration here. A "full-day trip" is defined as an excursion with at least four hours of collecting time, five to six hours of physical effort being, in our experience, the upper limit for a European in the Tropics. Half-day trips and those made under bad weather have been excluded. The distribution of full-day trips in the sense given above, by years, is as follows (number of days given in parentheses): before 1959 (21), 1959 (19[+9]), 1960 (16), 1961(27), 1962(11), 1963(44), 1964(22), 1965(11), 1966(32), Total (203[+9]). The card-file, however, contains all observations, including short or unsuccessful trips. Since April, 1961, the individual frequency of any species noted has been also added tc the statistics of species, making available presently the results of 138 full-day trips for this data.

Initially inspired by the above-cited papers, which deal only with the highest values observed, the author has here attempted to give a more complete treatment of the matter. Certainly, a greater number of observations would lead to more reliable conclusions; however,other duties may oblige the author to cease this very time-consuming statistical work. Thus this contribution is now presented, although it is certainly susceptible to improvement.

II. TYPES OF BUTTERFLY FREQUENCY

The total number of species and individuals found in a given day is, naturally, a complicated function of several independent frequency variables:

1. The individual frequency, i.e., the number of individuals

2                                           Ebert: Types of frequency

which characterizes a given population. This could also be called intensive frequency.

2.  The seasonal variation of the individual frequency, which could also be called seasonal frequency^

3.  The frequency of populations of a given species within a certain faunistical unit. This could be called also regional or ex-tensive frequency"!

In this contribution, the intensive and extensive frequency of species, as used during this study, will be discussed first. Then the number of species and individuals found during full-day excursions ("Daily frequency of species" and "Total frequency of individuals") will be discussed, with the seasonal variation of the last two categories. A number of more than 100 species found in one day is not rare in Brazil. The highest value obtained hitherto was 153, but as will be shown below, a value of 200 is well within real possibility2, at least for a group of three or more collectors.

III. THE REGIONAL (EXTENSIVE) FREQUENCY

Reliable information on the frequency of butterfly species within a given region is extremely difficult to obtain, especially when the number of species present in it is 600 or more, as is the normal case in the Neotropics. Some approximation is possible if the collector frequents a great number of localities and if the excursions are equally distributed through the year. Then the number of observations of any species recorded during a given period must be a relatively simple function of the three frequency types mentioned above, and the only one in which the extensive frequency plays an important part. For practical use, a period has been chosen during which the most frequent species (e.g. Pyrgus oileus oreuSj Euptyohia hermes^ Helioonius erato phyllis, etc.) were noted more than one hundred times. This period normally covered five years. As all notes on the card file were used, the observations made on days not counted as collecting days (half days and those in bad weather) are also included. The number of observations for the various species have been grouped into five classes:

Very isolated (ii) 1 or 2 times seen during period chosen Isolated                                  (i) 3 to 9 times "

Moderately distributed (id) 10 to 20 times " Well distributed (d) 21 to 50 times " " Very well distributed (dd) more than 50 times seen during period

chosen

2 After this was written, higher values (cf. footnote 4) were  obtained

by other collectors with more intensive methods (ten hours in   the

field—a physical exertion possible only to very well trained   persons, in a tropical climate).

Frequency of Butterflies in Eastern Brazil

3

Three regions were studied in this way: the eastern part of the State of Pernambuco, where the author collected from April 1957 until January 1962; the eastern part of the State of Sao Paulo [April 1962-December 1966]; and the surroundings of Rio de Janeiro (State of Guanabara) with the neighboring areas of Niteroi and the Organ Mountains (State of Rio de Janeiro; 1951-1956 and 1962-1966). The total numbers of species identified during this time are: 532 (Pernambuco), 742 (Rio) and 759 (S. Paulo). If the western parts of the State of Rio de Janeiro (Serra do Itatiaia) were to be included, the number for "Rio" would be 853. The number of species actually caught may be 10 to 20% higher, as a great number of species, mostly Hesperiinae and Theclinae, but also many Ewptyohia and some Rio-dinidae, are awaiting determination, still not being noted in the card file.

The statistics gave the following result:

Table 1: The number of species belonging to each of the five ranks of regional frequency and (in parentheses) its relative percentage.

Eastern Rio de Janeiro Eastern Median Rank Pernambuco _____etc._____ S. Paulo Percentage

247   (33)                      241    (32)                  32

228   (31)                      286   (38)                  36

I 10   (15)                        131    (17)                    15

I 17    (16)                             83    (I 1)                      13

40   (5)                        18   ( 2)                   4

742                                     759                              100

The median percentages in the last column have been calculated with consideration of the relative weight of each region, e.g., for the rank dd: (6 x 532 + 5 x 742 + 2 x 759) : 2033. A more detailed form of Table 1 is given in Appendix I, containing the values separated for all families (and subfamilies) for the Hesperiidae.

It may be noted that two-thirds of all species found have been seen less than 10 times, i.e., in less than 10% of the excursions made, and that only 4% of the total species have been seen in 50% of the trips.

There can be no question that the greater part of the neotropical species has a very scattered distribution, being encountered only occasionally. Populations must be generally of very restricted size and widely separated from one another. Such behavior would be easily understood in a land with great variability of biotopes. It is, however, difficult to understand in tropical Brazil with its very monotonous vegetation (rarely more than half a dozen natural ("climax") associations in areas of hundreds of square kilometers).

4

Ebert: Intensive frequency

IV. THE INDIVIDUAL (INTENSIVE) FREQUENCY

The following five-rank scale has been used by the writer, all ranks referring to the number of individuals observed (for the greater part not caught) of any species, during at least four hours of field work:

At the beginning of the statistical work, these ranks were checked by real counting of a limited number of species. After some training it proved easy to estimate the limiting values with sufficient reliability. Thus, the values used here are the result of estimation. They are adapted to the frequency features typical for the Neotropics; in other faunistical regions different limiting values may be more convenient, especially in the Holarctic region with its frequently high number of individuals in a limited space, such as in meadows.

Table 2 summarizes the results of the 138 full-day excursions, for which the statistics of individual frequency have been made. To permit comparison, all real values of the ranks have been transformed into percentages, and at the end of every column the number of species involved is given. The first value of every column indicates the median value of the respective group, the values in parentheses being the extreme percentages observed; for example, rr 53 (45-63) signifies that 53% of the observed species were in the rank rr (only 1 or 2 specimens seen during one day) and that the real values of percentage of rank rr varied between 45 and 63. The same holds good for the number of species given in the last line, e.t., 91 (57 -114) signifies that the number of species found in one day varied between 57 and 114, 91 being the median value of all days.

Table 2: Percentages of individual (intensive) frequencies resulting from T38 full-day trips in eastern Brazil.

Frequency of Butterflies in Eastern Brazil

5

Table 2 - continued

A) 24 excursions made in northeastern Brazil (states of Pernambuco and Paraiba) between April, 1961, and January, 1962.

B) 14 excursions made near Pocos de Caldas (Minas Gerais, 1250 m) from 27 January to 13 February, 1963.

C) 4 excursions from 1 to 5 May, 1963, at the same locality as B.

D) 6 excursions between 23 and 28 March, 1964, at the same locality as B.

E)  7 excursions to the same locality as B, between 10 and 18 December, 1966.

F) 17 excursions in mountains of southern Brazil (Organ Mts., Itatiaia Mts., Serra de Bocaina, Campos do Jordao; States of Rio de Janeiro and Sao Paulo), during January, February, March, July, September, November and December; elevation between 800 and 1700 m.

G) 40 excursions in medium altitude (500 to 700 m) forest areas of eastern Sao Paulo, distributed through all months of the year.

H) 16 excursions in the humid coastal region between Rio de Janeiro and Santos (below 200 m), from December to March and from July to August.

6

Ebert: Daily frequency

J) 10 excursions in the "Cerrado" of central Sao Paulo (700 to 800 m), in January, April, May, June, August, October and November.

The median values are calculated with consideration of the relative weight of the group values.

From these values it is clear that nearly half of all species are in the lowest rank (rr), only one or two specimens having been seen during a normal full-day excursion. In addition, more than 80% of all species were represented by less than ten individuals, whereas only 3% were seen to the extent of more than 20 and only 1% in more than 50 individuals. This phenomenon is quite contrary to that familiar to collectors in the Holarctic where a great number of individuals is typical for many species. Thus frequently a Neotropical forest or "Cerrado" appears to the collector surprisingly poor in butterflies. This scarcity of individuals of most of the species seen on the wing is, however, by and large compensated by the high number of species found. The median value of the 138 excursions, 77 species, is higher than the highest records found in the Holarctics. Thus it can be stated:

In eastern Brazil the individual frequency of butterflies is

tenerally very low\ The success of an excursion is exclusively etermined by the number of species found. The higher the number of species found during a trip, the higher the chance to find regionally (and/or individually) rare species (ranks ii and i of the foregoing section, rr and r of this section).

V. THE DAILY FREQUENCY OF SPECIES

The extreme values mentioned above (Table 2, G) of 15 and 153 species respectively for a single day are the extremes of all values noted by the author since July, 19583. The highest values, 150 and 153, were observed on two successive Sundays, the total number of species found during these two days being 204.

For statistic interpretation, the values of daily frequency of species have been grouped in the following classes: 16 to 20, 21 to 25, 26 to 30, 31 to 40 etc., until 151 to 155. Table 3 and Figure 1 show the frequency of these classes. The individual values are shown in Tables 4 and 5.

The special curves for Northeastern (N) and Southeastern (S) Brazil show, that the tendency for median values is more pronounced in the North, where the extremely low and high values found in the South are lacking. This may be the consequence of the more uniform climate during the year, i.e., the influence of a lower seasonal variation of the frequency.

■3--------------------------------------------------------------------------------

See, however, footnote 4.

Frequency of Butterflies in Eastern Brazil                       7

Table 3: Number of days (X) with values (Y) of daily frequency of species

10 16 21 26 31 36 4] 46 51 56 61 66 71 76 81 86 91

Y to to to to to to to to to to to to to to to to to

15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95

XN 000020 I I 11075511564

X$ I 0 2 0 0 6 2 4 8 II 12 12 ]j II 9 8 4

XT I 0 2 0 2 6 35 92I1917I6 22I4I4 8

96 101 106 III 116 121 126 131 136 141 146 151 No. of

X..: Northeastern Brazil (Pernambuco and Paraiba).

X'. Southern Brazil (Rio de Janeiro, Sao Paulo, Minas Gerais}.

X-p Total values.

10 20 30 40 50 60 70 80 90 100 110 120 130140 150 Species per day

Fig. 1 Daily frequency of butterflies in Eastern Brazil

N: Northeast (Pernambuco, Paraiba). SI Southeast (Rio de Janeiro, Sao Paulo, Minas Gerais). T: Total

8                                        Ebert: Variation of frequency

Hie curve of the total values shows a very pronounced subdivision into three parts:

28 days (14%) with fewer than 55 species in one day 123 days (60%) with 56 to 90 species 52 days (26%) with more than 90 species

The first group can be considered as corresponding to bad collecting days, the second as normal, the third as referring to very good days.

Theoretically, bad and fine days should be equally frequent, and the curve must be symmetrical. The asymmetry observed is purely subjective: the experienced collector naturally knows the season of the year which gives the best results (cf. the following section) and concentrates his efforts to it.

The median value of all 203 excursions is nearly the same as reported above (Table 2) for 138: 78 species per day. Thus a number of 75 to 80 species for one day may be considered as the normal value" in eastern Brazil.

VI. THE SEASONAL VARIATION OF THE DAILY FREQUENCY OF SPECIES

Very little is known, quantitatively, of seasonal variation of butterflies' frequency in the Neotropics, although any collector knows that during the dry season the number of species and individuals diminishes. Tables 4 and 5 and figures 2 and 3 will give some idea of this variation; they show the singular values already used for the construction of table 2 and figure 1.

Table 4: Seasonal variation of frequency of butterfly species in Northeastern Brazil (States of Pernambuco and Paraiba)

Reg ion_____Jan.______Feb.______March______Apr.______May______June

Frequency of Butterflies in Eastern Brazil 9 Table 4 - continued

Region July______Aug.______Sept.______Oct._____Nov.______Dec.

CR Coastal region (sea level to 100 m); 52 excursions

M           Median values for the coastal region

A "Agreste" (higher and less humid elevations at some

distance from the coast; 300 to 900 m); 16 excursions

M„ Median values for the "Agreste"

The coastal region was originally covered by rain forest, presently substituted for the most part by sugar cane plantations. The 'Agreste" receives less precipitation than the coastal zone, but still sufficient to allow the growth of a low forest, partly defoliated during the dry season.

Table 4 and figure 2 show the following: in the coastal region the seasonal variation of frequency of species is low. Its maximum value coincides with the period of maximal precipitation. With the beginning of the dry season, the frequency diminishes only slowly, not accompanying the decrease of rain frequency. Apparently the densely foliated forest conserves appreciable humidity during the dry season. The lowest value, which is still higher than half the maximum value, is reached at the beginning of the rainy season. The secondary peak at the time of greatest drought may be caused by the appearance of a second generation of many species.

In the "Agreste" the precipitation is much lower than near the coast, however its periodicity is the same. Also here the butterfly frequency reaches its lowest value at the beginning of the rainy season, and the diminution of frequency is also slow after the end of the rains, showing the same secondary peak during the strongest drought. The number of species is lower in all months than near the coast, certainly a consequency of the impoverishment of the flora.

10

Ebert: Variation of frequency

Species per day

WO n

I E M IYYYLWYEIKXMXR Months

Fig. 2 Seasonal variation of the daily frequency of species in

Northeastern Brazil, compared with the monthly distribution of rainfall.

Frequency (scale to the left): F

CR

Rainfall (scale to the right): R

R

Coastal Region (sea level] "Agreste" (300 to 900 m) Recife (Pern.; sea level) Garanhuns (Pern.; 850 m)

Interpolated parts of the curves are broken.

The minimal values are lower than half of the maximal. This greater seasonal variation is similar to that we shall find in the south (figure 3). The faunistic character reflects the same analogy: in the "Agreste" nearly all hylaean species are lacking, being partially substituted by typical species of the south.

Figure 3 and Table 5 show the seasonal variation in southern Brazil. Here the collecting localities are subdivided, in accordance with their floristic-faunistic character, into three groups, low regions (below 200 m) (L), higher regions (500-1500 m) (H) and savanna-like "Cerrado" (C).

The number of observations in the lowlands and in the "Cerrado" is still low, and the values given here may not be fully representative. In the coastal zone the highest values have been observed in April and May, as the author remembers from the years 1951 to 1956 when he was not yet noting frequency data. This high season may continue until July (verbal communication by K. Brown).

Frequency of Butterflies in Eastern Brazil n

Table 5 Seasonal variation of the daily frequency of butterfly

C 36                                                          90 62 84

68 ___________________________________________Q[__________

M                       36                                                           90 704 84

Region July Aug. Sept. Oct. Nov. Dec.

68 82 55                                                         37

L 75                                      59                                                         55

75                                      74                                                         62

86                                      76

_____86_____________________________

ML                       78 82 70                                                         51

23                 39 15 68

37                 • 55 40                                      64

62

12                                    Ebert: Variation of frequency

Table 5 - continued

Region______July______Aug.______Sept.______Oct.______Nov.______Dec.

MH 41 39 45 68 58 73

C                                          77                                        39 2]

________________________________________49____________

Mc                                        77                                        39 35

L Low regions near the coast with nearly tropical conditions, between Rio de Janeiro and Santos (18 excursions).

H Higher regions (500 to 1500 m) in the Interior, less humid and

with a tendency to subtropical conditions, however, sufficiently humid to allow forest vegetation (Pocos de Caldas, M. G.; eastern S. Paulo; Itatiaia Mts., R.J.' some data are from the Serra do Mar); 103 excursions.

C Savanna-like "Cerrado" with only very narrow strips of low wood along the water-courses, in the Interior of the State of Sao Paulo (10 excursions).

ML MH MC M©dian values for the zones L, H, and C.

The dependence of the frequency of species on climatic conditions is evident from figure 3. In the humid coastal zone the seasonal variation is small, the lowest values corresponding to the beginning of the rainy season. The same has been observed in tropical northern Brazil (cf. figure 2). Perhaps this feature may be typical for other Neotropical areas. According to a note of D. Zayziw (1958:5-7), the behavior of Coleoptera is quite different: the Cerambycidae studied by him showed clearly their maximum frequency at the beginning of the wet season (October-November). This may be the result of the different way of life of the larva and of different "hibernation" stage; as egg or imago in butterflies, as larva or pupa in beetles. Butterflies reach their maximal frequency at the end of the wet season.

the "Cerrado" of ltirapina (S.P.) by the author's son Karl and Dr. Keith Brown of Rio de Janeiro, on May 21, 1967. The value transforms the above median value for May to 93. This value has been used for the construction of Figure 3, but was relegated otherwise to avoid tedious rewriting, recalculation of tables and reconstruction of curves. Still higher values were found by Dr. Brown, made by more intensive collecting (cf. footnote 2) in Central Minas Gerais and near Rio de Janeiro, including several days with counts of over 200 species and one of more than 250 (Brown, verbal communication).

Frequency of Butterflies in Eastern Brazil 13

Species per day 120-1

Species per day r120

Month I n

H M II IX

zr xn

300 250 200 150-1 100 J 50 J

LUb

^

300 250 200 1 50 MOO 50

Month I nm lYlIMKISXII

Fig. 3 Seasonal variation of the daily frequency of species in

Southeastern Brazil, compared with the monthly distribution of rainfall

Frequency (upper diagram; details cf. explication of Table 4): L: Lowlands (below 200 m). H:

Highlands (above 500 m). C: "Cer-rado"

Rainfall (lower diagram): Ub Ubatuba (S. Paulo; sea level)

R. CI. Rio Claro (S. Paulo; 600 m; at the limits of forest zone and "Cerrado").

Interpolated parts of the curves are broken.

In the less humid Interior, the type of variation is similar: highest frequency at the time of the last rains, lowest values at the end of the dry season. The decline of the curve, at the beginning of the dry season, is sharper and the seasonal variation greater than near the coast.

In the forests of the Interior the curve is simple: a high peak in April--May with more than 100 species every day, and a notable decrease in June and July. The minimum with less than fifty species

14

Ebert: Daily frequency

occurs at the end of the dry season; the rise to the new maximum is slow.

In t^ie "Cerrado" the general trend of the frequency curve is more similar to that of the coastal zone, notwithstanding the different character of the "winter" with its hot days and cold nights. The minimum occurs also at the beginning of the wet season here. It has been a great surprise to the author to observe during the "winter" (June - August), a much higher number of species, in the midst of a totally dry vegetation, than at the beginning of the wet season (October - November), when all the vegetation is covered with fresh foliage and many blossoms.

Generally the best time for collecting in southern Brazil is the end of the wet season, April - May. At this time a daily catch of more than a hundred species can be expected in a good locality, with some variation in the vegetation and many blossoms.

VII. THE DAILY FREQUENCY OF THE TOTAL NUMBER OF INDIVIDUALS

It is possible to reach some quantitative conclusions on the total number of butterfly individuals seen during one day, including all species identified, with the aid of the values of the daily frequency of species and the individual (intensive) frequency of these species grouped in the five ranks discussed above. Thus 1.5 is the median value for the rank rr, 6 for the rank r, etc., (rr) is the number of species found in rank rr, (r) the same for rank r, etc. The following formula for the total number of individuals results:

FTI = 1.5 (rr) + 6 (r) + 15 (rf) + 35 (f) + 100 (ff)

Such a calculation is, naturally, quite schematic and only an approximation to the real number of the individuals on the wing, but it furnishes comparable values for the different days and regions.

The following table and Figure 4 represent the statistics of FTT-values for the same 138 excursions used in Table 2. The group here includes hundreds of individuals (the individual values are shown in Table 6):

301

to

400

22

1201

to 1300

2

401

to

500

22

1301

to 1400

2

501

to

600

27

1401

to 1500

2

601

to

700

8

1501

to 1600

0

701

to

800

7

1601

to 1700

1

801

to

900

9

Frequency of Butterflies in Eastern Brazil 15

200 400 600 800 1000 1200 1400 1600 Individuals per day

Fig. 4 Variation of the number of individuals of butterflies in Eastern Brazil.

Northeastern and southeastern Brazil are not separated in Figure 4, because of the low number of values (24) from the northeast. Figure 4 shows the same subdivision into three groups as the sum curve of Figure 1:

9 low days (7%) with less than 201 individuals per day 92 "normal" days (66%) with 201 to 600 individuals per day 37 high days (27%) with more than 600

The motive for the greater frequency of high days in relation to low ones is the same already discussed in connection with Table 2.

VIII. THE SEASONAL VARIATION OF THE TOTAL NUMBER OF INDIVIDUALS

In Table 6 and Figure 5, the values are grouped in accordance with the months and with the regions, in analogy with Table 4 and 5 and Figures 2 and 3.

Comparison of Figure 5 with Figures 3 and 4 shows that the seasonal variation of number of individuals is the same as that of species, with the maximum and minimum parts of the curves more pronounced. The upper diagram (I) represents observations in tropic climate: in the northeast and at the Littoral between Rio de Janeiro and Santos. The lower diagram (II) shows the situation in the less humid higher regions of southern Brazil. All frequency curves show more than one maximum and interesting relations with the rainfall curve.

16                                     Ebert: Frequency of individuals

Table 6 Seasonal variation of number of individuals of butter-

Mu 483 568 521 907 984 459

H

Region July Aug. Sept. Oct. Nov. Dec.

NF 456 249 354 542 438 337

873

MNE 652 317 442 542 650 337

198 77 354

Frequency of Butterflies in Eastern Brazil 17 Table 6 - continued

Region______July______Aug.______Sept.______Oct.______Nov.______Dec.

M„

479 509 560 59]

528

124 168 360

517 515

515 231

]<

215

245 376 442 549 613

445

54 294 334

580

189 229 268

229

227

231

227

580

344

372

NE C L H

Group A of Table 2 Group I of Table 2 Group H of Table 2 Groups B to G of Table 2 etc.: Monthly median values of the four groups.

The main maximum coincides in the northeast (F^) with the rain maximum: in the Littoral (Fj) it is retarded two months, and in the higher forest areas (FJ four months in relation to the rain maximum. Two secondary maxima appear in the northeast during the dry season; probably only one secondary maximum exists here, the minimum value for December not being representative. In the coastal area of the south the minimum coincides with the beginning of the rainy season, being retarded four months against the rain minimum. The second minimum of the FT curve may be fictitious, the only value for February not being representative. The same doubt exists for the secondary maximum of the FH curve, as the single value for October may be exceptionally high. Typical for the semi-dry forests of the Interior is the sudden decline between May and July, at the beginning of the strongest drought. It may be the effect of the first cold nights. The minimum corresponds to the end of the dry season.

In the "Cerrado" the main maximum coincides with that of the other regions of the South, the minimum with that of the Littoral, being retarded four to five months after the driest months. The secondary maximum in the month of August is still dubious, as the only high value perhaps is not representative.

18

Ebert: Frequency of individuals

Fig. 5 Seasonal variation of frequency of individuals in Eastern Brazil

Upper diagram (I): Areas with tropical features

Frequency (scale to the left): FNE: Brazilian Northeast.

F, : Littoral between Rio de Janeiro and Santos.

Rainfall (scale to the right): RR : Recife (Pern.).

Ry : Ubatuba (S. Paulo)

Lower diagram(II): Regions with the beginning of subtropical features

Frequency (scale to the left): FH: Highland forests.

FQ: "Cerrado"

Rainfall (scale to the right): RRC: Rio Claro (S. Paulo; 600 m; at

the limits of the forest zone and the "Cerrado").

Frequency of Butterflies in Eastern Brazil 19

Individuals per day 8OO-1

Rainfall I                      mm

i n E^^w^Y^m^m^m^K^T^.

20

Ebert: Conclusions

A great variation of frequency within the same month is notable, e.g. for the F values of 243-872 (January), 339-1249 (February), 351-1460 (ApriT), 298-1760 (May), 189-865 (December). Here the influence of the annual variation may be reflected. Also in the Neotropics "good" and "bad" years exist. Some values typical for this type of variation are given below in Section Xc.

IX. CONCLUSIONS From the foregoing data the following conclusions may be drawn:

(a)  Generally, butterflies appear in the Neotropics in very low numbers of individuals.

(b)  This scarcity refers to the low density of populations as well as to the low numbers of individuals belonging to many of them.

Thus the frequently defended hypothesis that any butterfly species may be abundant at one, possibly hidden, locality and during a defined, probably very short, time seems to be not valid for eastern Brazil. Here certain species have been found only once in only one specimen in ±450 excursions during 15 years at the most varied localities between the 6th and the 23rd parallel.

(c)  Only very few species are seen everywhere and always   in great number. These are common and widely distributed species  as

Agraulis vanillae^ Dryas julia^ Pyrgus oileus orcus3 Euptychia  hevmes3 Eelioonius erato phyllis^ Phyoiodes olaudinay etc.

(d)  The seasonal frequency variation of species as well as of individuals is very large, more than would be expected in tropical zones. The variation in the number of individuals is still stronger than that of species.

(e)  In zones near the equator the frequency variation accompanies the rainfall curve, showing a second peak during the dry season.

(f)  In the southern areas, partially with subtropical tendency, the frequency variation is much retarded in relation to the rainfall curve. Temperature may be of influence at the end of the summer, when the first cold nights coincide with an abrupt diminution of individuals.

(g)  A second, much lower, frequency peak may exist two months                           * before the highest peak of the rainy season (curves FT and Ff in

figure 5); this feature needs confirmation, as the number of observations is still insufficient.

(h) The number of individuals seen on the wing in the Neotropical Region may be more or less equal to that seen on a summer day in the Holarctic. However, the relation between number of species and their individuals is the inverse, the number of species compensating their lower number of individuals.

Frequency of Butterflies in Eastern Brazil 21

(i) A catch of more than 100 species per day is normal during the short time of maximal frequency. The highest value, hitherto found by the author, has been 162, other collectors being still more fortunate (cf. footnote 4),

The median values of individuals per day are: 518 in the northeast, 461 in the coastal zone of southern Brazil and 562 in the higher forest regions of the South, the total median value of the forest zones being 542. Since we found as a median number of species a value of 77 to 78 for a day, the median number of individuals per species per day is only seven, a very low value compared with that familiar for the collector in the Holarctics (cf. Newcomer, 1962). In the "Cerrado" the median number of individuals is lower: 350. The median value for all 138 excursions is 528.

Thus we come to the following conclusion: in the Neotropic the relation between the number of species and their number of individuals is the inverse of that normally observed in the Holarctic; many species with very low number of individuals in the Neotropics; far fewer species in far more individuals in the Holarctic. The total number of individuals observed during one day may not be very different in both regions.

X. ADDITIONAL OBSERVATIONS

In the following we shall discuss briefly the influence of the number of observers and of the time of observation.

(a) Influence of the number of observers

For the study of the influence of the number of hunters on the number of species found, sometimes a separate analysis has been made for the catch of the present writer and of that of his son (presently 19 years old); the results are given in Table 7 (the number in parentheses refer to percentages):

Table 7 Number of species found by two hunters (with percentages)

22

Ebert: Influence of collecting hours

Date 1963 2/1 I I

1963 4/1 I I

1963 10/1 I I

1966 6/1

1966 29/V

1966 9/V]

19/VI

1966 I5/VI1

Father 49(42) 47(50) 55(44) 22(23) 37(38) 26(31) 38(35) 29(38) Son 32(27)21(23) 30(24) 38(39) 26(27) 38(45) 33(31) 27(35) Both 36(31)25(27) 41(32) 37(38) 34(35) 20(24) 37(34) 21(27)

17

93

126

97

97

84

107

77

27/1   62: Recife (Pern., sea level)

27/1   63: Pocos de Caldas (Min. Ger., 1300 m)

2/XI   62 to I6/III 63: Rio Claro (Sao Paulo, 600 m)

6/1,   29/V and 19/VI 66: Araras (Sao Paulo, 600 m)

9/V I and 15/VIII 66: Itirapina (S. Paulo, 750 m)

The median and extreme percentage values are: Found by the first hunter (father) only: 38.8$ (23

(Forest)

("Cerrado")

Found by the second hunter (son) only: Observation common to both hunters:

50) 30.6/o (21 - 45) 30.6/o (21 - 46)

During the 15 full-day trips on which these separate annotations were made, an average of only 31% of all species found, i.e., less than one-third, have been collected by both hunters, and more than two-thirds only by one or the other. It is evident that by an increase of the number of hunters the number of species found would increase also. The number of 150 and 153 species caught by two hunters on the 8th and 15th of May, 1966, and the observation that on these two succeeding Sundays the total number was 204, only 98 (48%) of them found on both days permit the conclusion that a team of three or four hunters, during the best season, may get easily more than 200 and perhaps 300 species during one day in eastern Brazil(cf. footnote 4), many more than Shull (1962) refers to for a trip of seven collectors in India. The frequently defended hypothesis that the Neotropical Region is the richest of all may thus be considered proven quantitatively.

(b) Influence of amount of collecting hours

The last example already suggests that for a series of succeeding days, every day species previously not seen, will appear, whereas others will seem to have disappeared. Really they all have been on the wing all the days, however most species are seen only occasionally owing to their very great scarcity. This observation corroborates the conclusion already drawn from other data: the majority of Neotropic butterflies are rare in the sense of extensive distribution as well as of intensive frequency. Thus some observations on the number of species found during a group of days may be of general interest. Table 8 summarizes the results of four stays in Pocos de Caldas (State of Minas Gerais; 1000 to 1500 m) during January, February and May of 1963, March of 1964 and December of 1966.

Frequency of Butterflies in Eastern Brazil                          23

X is the number of species found on the single days, Y the number of species found for the first time, during the respective stay, on every day.

Table 8 Increase in number of species during collecting periods

We note that every new day brings species not seen before, where others seem to disappear. Thus the total number of species found during a several day period is two to three times higher than the median species number per day.

But also any subsequent visit reveals species never seen during the earlier, as is shown in the following table, which includes more recent observations:

Period Jan./Feb. 1963 (14 days) May 1963 (4 days) March 1964 (6 days) December 1966 (7 days) ApriI/May 1967 (6 days)

A list of these 572 species is given in the Appendix

A catch of 572 species during five stays with 37 collecting days during six months may be unrivaled outside of the Neotropics. Diagram I of Figure 6 represents as accumulative curve the continuous increase of species with every new stay; the curve rises steadily without any sign of asymptotic approximation to a final value. There

24

Ebert: Accumulative values

is a vague possibility only to estimate the total value of species occurring at this locality (700 to 800?).

It is highly interesting that the same feature holds for longer periods of time, as tables 9 and 10 and the diagrams II, III, IV of figure 6 show.

Table 9 First observation of species during five successive years in Eastern Pernambuco (accumulative values in parentheses)

Table 10 First observation of species during successive years in Southeastern Brazil

5                                                              5

Year 1951 1952 1953 1954/5 1956 1957 1963-1965 1966 168 85 50 40 43 22 42

460 545 595 635 678 700 742

4

Rio de Janeiro,etc.

In some cases cases two or three years have been combined to one observation period, to warrant equal weight of the periods (+_ equal number of excursions).

Spe- 292 cies

Accumulative 292 Values

Frequency of Butterflies in Eastern Brazil                        25

Table 10 » continued

Year______________1962 1963 1964-1965 [966

Species 332 277                   83 67

Accumulative 332 609                 692 759 Values

Eastern Sao     Paulo

All curves show, for three long-period observations, the same nearly rectilinear rise of the curves as that of the short-period observations of Pocos de Caldas. The number of species is higher, owing to the greater number of observation days (more than 100), but the general form of the curves is the same. There is not the slightest sign of asymptotic approximation to a final value. The real number of species remains unknown although in two areas already values of +750 species have been reached. To conclude from the material found Tn other collections and by ourselves in other parts of the states of Rio de Janeiro and Sao Paulo, in both states the total number of species surely surpasses 1,000, perhaps reaching 1100 or 1200.

Table 9 shows the different behavior of different families; the frequent and easily noted groups such as Brassolidae, Danaidae, Heliconidae, Pyrrhopyginae, Papilionidae, do not contribute to the increase of number of species in the later years. This increase is due essentially to the "little species" as Hesperiidae, Lycaenidae and Riodinidae which constitute +_ 2/3 of the total number of species (cf. Appendix I). Collectors who do not give special attention to these groups will never know their local fauna in the Neo-tropics. All our observations on frequency and distribution of species refer thus essentially to these three families.

The fact that at frequently visited localities species never seen before appear constantly, whereas others seem to disappear for a very long time, and the observations made on frequency above may be interpreted as follows:

(1) In the (eastern) Neotropics the populations of many species of butterflies, especially of the most numerous families Hesperiidae, Lycaenidae and Riodinidae, are small (intensive frequency low).

(2)  The different populations of such species are dispersed widely and with great distances between them (extensive frequency low).

(3)  These populations do not maintain constant their localities of habitat but migrate continuously within a great area of favorable biotopes leaving their old habitats and creating new colonies which will be left after a short time too. Such behavior would explain the continuous appearing and disappearing of species. Perhaps it is typical for many Neotropical creatures, as it is well known also in birds and in the primitive men of the forest, the Amerindians.

26

Ebert: Accumulative values

Number of species 1000i

n

Stay Ibl Iu mu n?Th Yrhyear1957 58 59 60 61

Pocos de Caldas [Mm.Ger.]

pernambuco

Number oF species -1000

L800

19 51 52 53 54/5 56 57 Rio de Janeiro

63 66

65

1962 63 64/5 66 Sao Paulo

Fig. 6 The rise of the number of butterflies species with prolonged observation time, in four selected areas. Accumulative curves. No asymptotic approximation to a final value has appeared!

Frequency of Butterflies in Eastern Brazil                          27

(c) Observations on annual frequency of species numbers

As the time-consuming statistical work used for this paper cannot be continued, in consequence of professional duties, some observations on annual variations may be added:

Tiuma (Pern.) 20.VII.58     26.VII.58     31.VII.58     3.VIII.58     9.VIII.58

102 116 99 101 129

" " 27.VII.59     8.VII.60     14.VIM.60     8.VIII.61     4.X.58

94 66 89 64 68

18.X.58 26.IX.59 8.IX.60 89 65 80

Camaragibe 7.IX.58 7.IX.59 24.IX.60 2.IX.61 (Pern.)                        93 104 83 87

From the above values it may be concluded that in really tropical areas the frequency variation between different years is low. Thus the seasonal frequency curves for such areas shown above (Table 4 and Figure 2) may be reliable, even though having only a limited number of observation years.

Foot of the Organ Mountains (R. Jan)

Litoral Norte (S.P.)

Itatiala South (R. Jan.)

The humid regions of southern Brazil seem to possess the same low annual variation of frequency as do the tropical zones.

Itirapina - 21.IV.63 3.V.64 I.V.65 1.V.66 6.VI.66 21.V.67 Brotas (S.P.) 90 62 68 81 84 162

13.VI.68 48

Araras (S.P.) 17.IV.66 24.IV.66 8.V.66 I5.V.66 19.VI.66 124 107 150 153 108

20.V.67 3.V.68 152 87

Mirassol (S.P.) 13.V.67 25.IV.68 109 68

28

Ebert: Annual variation of frequency

Faz. S. Jose, 13.IV.63 23.V.63 5.V.64 28.111.65 I6.V.65 I9.V.66 Rio Claro (S.P.) 143 108 75 82 84 107

In the less humid areas of the interior, of nearly subtropical climate, the annual variation of frequency is great, as is the seasonal. The years 1963, 1966 and 1967 were good years; 1964 and 1965 average and 1968 a poor year. The extreme high values of daily frequency mentioned above will be reached only in exceptionally good years. Particularly striking is the difference between 1967 and 1968, notwithstanding the lack of clear cliniatological difference which could be observed between the rainy seasons of 1966-67 and 1967-68.

(d) Observations in the high Tropics

From high tropical regions only data for the daily frequency of nine days are available: 4.-9.XI. British Guiana (Parika, Atkinson Field), 12.XI. Belem (Utinga), 14-23.XI. Rio Ampari (Amapa; Serra do Navio and Porto Platon).

1959 4/XI 7/XI 8/XI 9/XI 12/XI 14/XI 19/XI 21/XI 23/XI

Species 71 69 57 81 86 51 62 60 66 per day

The median value (69) is lower than that from eastern Brazil (77-78), and also the total number for the four days in Amapa (179) is lower than comparable values from Pocos de Caldas, but as the observations are from one month only, the values may not be representative for the year.

The following table shows the percentage by which the different families are represented:

Frequency of Butterflies in Eastern Brazil

29

With respect to the relative frequency of families, the coastal zone of British Guianas shows the same predominance of the Hesperiidae as eastern Brazil, whereas at the borders of the Amazon the Riodinidae are the most frequent family. Perhaps during another time of the year the proportions may be different.

ACKNOWLEDGMENTS

A great number of the entomological observations used in this paper were collected at the occasion of geological excursions made with the aid of the "Fundacao de Amparo a Pesquisa do Estado de Sao Paulo" and the "Conselho Nacional de Pesquisas" to whom the author is deeply obliged. A very substantial part of the material here used was contributed by the zeal and skill of the author's son Karl Rudolf to whom he owes cordial thanks. Dr. K. Brown (Rio de Janeiro) kindly revised the manuscript and gave valuable suggestions to its emendation.

30                                                 Ebert: References

LITERATURE CITED

BROWN, K. S. AND 0. H. H. MIELKE

1967     Lepidoptera of the Central Brazil Plateau I. J. Lepid. Soc, 21: 77-106, 145-168.

1968     Lepidoptera of the Central Brazil Plateau III. J. Lepid. Soc, 22: 147-157.

HEMMING, F.

1955 My highest catch of butterfly species in a single day: Digne (France), 6th August 1926. Lepid. News, 9: 144-145.

LANGER, T. W.

1955 One day's species in the French Alps. Lepid. News, 9: 203-204.

NEWCOMER, E. J.

1962 Collecting is still good in the Northeast. J. Lepid. Soc, 16: 67-70.

REMINGTON, C. L.

1955     How many butterfly species in one day. Lepid. News, 9: 77-78.

SHULL, E. M.

1958 My highest catch of butterfly species in a single day

(4th June 1961) at Mussoorie, India. J. Lepid. Soc

16: 143-145. 1962 Over one hundred butterfly species caught in a single

day (3rd June 1961) at Mussoorie, India. J. Lepid. Soc.

16: 143-145.

WILTSHIRE, E. P.

1956     My highest butterfly catch of a single day in the Middle East. Lepid. News, 10: 116-118.

ZAYZIW, D

1958 Fauna do Distrito Federal, XVIII: Contribuicao para o estudo dos Longicorneus do Rio de Janeiro (Coleoptera-Cerambycidae). Bol. Mus. Nac, N.S., Zool., 189: 26 p., 2 fig.

Frequency of Butterflies in Eastern Brazil                         31

APPENDIX I

Extensive frequency of the different families of butterflies in three selected areas of Eastern Brazil (Values in parentheses refer to percentages.

(A) Eastern Pernambuco

Total 76 (14.3) 62 (11.6) 2 (0.4) 54 (10.) 15 (2.8)

194 (36.3)

32

Ebert: Distribution of frequency ranks

(B) Rio de Janeiro and surroundings Hesperi idae

Itho-

Rank Hesperiinae Pyrginae Pyrrhopyginae Urbaninae _________________________1 a___ __v_________________mi i dae

21 (37)        6    (25)

2] (37)      I]    (40)

7 (12)        3    (12)

7 (12)        4   (15)

I (2)           2   (8)

Total 137 (18.4) 84 (]] .3) 5 (0.7) 57 (7.7)   26   (3.5)

283 (38.2)

(C) Eastern Sao Paulo

Hesperi i dae

Itho-

Rank          Hesperiinae       Pyrginae        Pyrrhopyginae       Urbaninae ...

ii                    42 (36)               27 (3D 9 (64)                    22 (41) 12 (38)

i                   49 (42)               33 (37) 5 (36)                    19 (35) 6 (19)

id                  18 (16)               18 (20) 0 -                        6 (ID 4 (12)

d                   7 (6)                10 (ID 0 -                       7 (13) 6 (19)

dd                   0 -                   I (I) 0 -                       o-4 (12)

i i                       0      -                    40 (39)                I    (33)                     14    (22)

i                      1    (100)             34 (34)               1    (33)                    19   (29)

id                     0     - 14 (14)               0     -                          8   (12)

d                     0     - 12 (12)               0     -                        16   (25)

dd                     0     - 1(1)                  1    (34)                      8   (12)

ToTal               1    (0.2)           101 (19.0)          3   (0.6)                 65   (12.2)

(B) Rio de   Janeiro and surroundings

Rank          Libytheidae        Lycaenidae Morphidae Nymphalidae

i i                       0      -                    46 (38)                 I    (17)                     18    (20)

i                      I    (100)             31 (26)               2   (32)                   28   (31)

id                    0     - 15 (12)               I    (17)                   10   (ID

d                   0     -                 21 (18)              I   (17)                 24   (27)

dd                     0     - 7 (6)                 I    (17)                    10   (II)

Total               1    (0.1)           120 (16.I)          6   (0.8)                 90   (12.)

Frequency of Butterflies in Eastern Brazil                         33

(C)     Eastern Sao Paulo

Rank______Libytheidae         Lycaenidae Morphidae         Nymphalidae

ii 0 -                     39 (43) 3 (43)                    20 (19)

i 0 -                     31 (34) 3 (43)                   32 (30)

id I (100)                15 (16) 0 -                       32 (30)

d 0 -                       6 (7) 0 -                        17 (16)

dd 0 -                       0 - I (14)                     4 (5)

Total I (0.1)               91 (12.0) 7 (0.9)                105 (13.9)

(A) Eastern Pernambuco

34

Ebert: Discussion of frequency ranks

DISCUSSION OF APPENDIX I

The detailed table of extensive frequency leads to the following conclusions:

(1)   In eastern Brazil, the total number of species is higher in the south than in the north, notwithstanding the more tropical character of the latter. Apparently the fauna of the northern parts (Pernambuco, etc.) is impoverished, the direction of the immigration here being from the south (Bahia) to the north.

(2)  Notwithstanding the different absolute values of the number of species, the relative frequency of the families is nearly the same in the three regions. Note for example the values for the Hesperiidae (36.3 - 38.2 - 35.81), Pieridae (5.1 - 5.1 - 5.8%), Nymphalidae (12.2-12.0 - 13.9%), etc. Remarkable are the low values for the Acraeidae, Brassolidae and Papilionidae in the northeast.

(3)  The percentage of the different ranks in the three regions are very similar: 28-38-15-13-6 in the northeast, 33-31-15-16-5

at Rio, 32-38-17-11-2 in Sao Paulo. Apparently the number of observations is high enough, and our values can be considered representative. Nearly all families show the typical predominance of the lower ranks, but there are some exceptions: in the Papilionidae the ranks are nearly equally distributed, and in the Heliconiidae frequent species prevail.

NOTE I: It must be kept in mind that the values of the Table refer to determined species only; the real numbers of species of Hesperiinae and Lycaenidae, and in a lesser degree those of Riodinidae and Satyri-dae, are much higher as a great percentage of these groups could not be determined,hitherto (some of them will prove to be new).

NOTE II: The subdivision of the Rhopalocera into Families and Sub-families, and the generic and specific names used here and in Appendix II are not in all cases in accordance with the use of other authors (d'Almeida, Clench, Emsley, Forbes, Fox, etc.). These modifications are based on taxonomic studies which are beyond the limits of problems discussed in this paper. Some remarks are made in Appendix II. To avoid phylogenetical speculations, alphabetical order has been chosen for the sequence of names.

Frequency of Butterflies in Eastern Brazil

35

APPENDIX II

List of the 572 species of butterflies found during 37 collecting days between December and May near Pocos de Caldas (State of Minas Gerais;

1000'to 1500 m)

We add a list of all species found at this locality as a contribution to the knowledge of the fauna of the transitional zone between the central highlands (Brown § Mielke, 1967, 1968) and the coastal zone of median Brazil.

Collecting places

R Remnants of high (primary) forest ("Mata") by the riverside

of the Rio Pardo (1000 m). F Low (primary?) forest of the mountains at the north of the

town ("Caixa d'Agua," 1300 m; highway to Botelhos, 1400 m). C Low (secondary ) forest ("Capoeira") near Cascata (1300 m)

and above the gorge of the Rio das Antas (1200 m). 0 Open grassland ("Campo") at the south of the town (1300 m)

and above the mountain-forest ("Serra de Pocos"; 1450 to

1500 m).

1, 2 .... 12: Months of observation

rr, r, rf, f, ff: Frequency, as discussed in the text.

Some observations made during shorter stays (August, October, November) have been included.

Acraeidae (7 sp.)

Actinote alalia (Felder, 1860) F, C, - 4, 12 - rr, r

pyrrha(Fabricius,1775)(=brasiliensis Almeida, 1922)

R,F,C, - 4,5,12 - r-ff carycina Jordan, 1913 R,F,C, - 3,5,12 - rr-f conspicua Jordan, 1913 F - 12 r-f melanisans Oberthur, 1917 (=rhodope Almeida, 1922)

R,F,C, - 2-5, 11-12 - rr-f parapheles Jordan, 1913 F - 12 - rr surima (Schaus, 1902) F,C, - 1,5,12 - rr,r

Brassolidae (7 spp.)

Blepolenis batea batea (Hubner, 1821) F,C, - 2,3, - rr-rf Caligo arisbe Hubner, 1822 F,C, - 1-4 - rr-r

illioneus illioneus (Cramer, 1775) R - 4 - rr Dasyophthalma rusina (Godart, 1821) F - 1-3,5 - rr Eryphanis reevesi (Doubleday § Hewitson, 1849) F - 12 - rr-r Opoptera aorsa (Godart, 1821) F - 3,12 - rr-r

syme (Hubner, 1821) F,C - 1-3,5 - rr-r

36                              Ebert: Butterflies of Pocos de Caldas

>

Danaidae (4 spp.)

Anosia gilippus gilippus (Cramer, 1775) R,F,C,0 - 2-5, 10-12 -

rr-r Danaus plexippus erippus (Cramer, 1775) F,C - 1-3,5,10 - rr-rf Ituna ilione (Cramer, 1775) F,C, - 1,2,5,11 - rr,r Lycorea ceres halia (Hiibner, 1825) F - 2,3 - rr

Heliconiidae (9 spp.)

Dryadinae

Agraulis vanillae maculosa (Stichel, 1907) F,C, - 1-5,8,10,12 -

rr-rf Dione moneta moneta Hiibner, 1825 R,F,C, - 5 - r,rf Dryadula phaetusa (Linne, 1758) F,C - 2-5 - rr,r Dryas julia julia (Fabricius, 1775) R,F,C, - 1-5 - rr-rf Philaethria Wernickei (Rober, 1906) F - 2,3,5 - rr,r

Hel i corn'inae (4 spp.)

Heliconius besckei Menetries, 1857 R,C,F - 1-5,11,12 - rr-rf

ethilla Latr. narcaea Latreille, 1820 R,F,C - 1-5,8,

11,12 - rr-rf erato phyllis (Fabricius, 1793) R,F,C,0 - 1-5,8,11,12

rr-f Eueides aliphera aliphera (Latreille, 1820) F,R,C,0 - 1,2,4,5 -

rr-rf

Hesperiidae (223 spp.)

Hesperiinae (113 spp.)

Anthoptus epictetus(Fabricius, 1793) R,F,C, - 1-5,11,12 - rr,r Artines aquilina (Plotz, 1883) F - 2 rr Callimormus beda (Plotz, 1886) R,F,C, - 2-5,12 - rr

interpunctatus (Plotz, 1884) R,F, - 2-5,12 - rr,r saturnus (Herrich-Schaeffer, 1864) C^5-r Cantha honor Evans, 1955 F,0 - 1-3,11 - rr Conga urqua (Schaus, 1902) F - 12 - rr Copaeodes jean favor Evans, 1955 F,C,0 - 2-5,12 - rr,r Corticea noctis (Plotz, 1888) F - 1,3 - rr Cumbre cumbre (Schaus, 1902) R,F,C, - 1-5,12 - rr-rf Cymaenes alumna (Butler, 1877) R,C, - 5,12 - r

tripuncta theogenis (Capronnier, 1874) 0 « 2 - rr tripunctata(Latreille, 1824) R,F,C,0 - 1-5, 10-12- rr-f Cynea corope (Herrich-Schaeffer, 1869) F-2-rr Dion meda (Hewitson, 1877) F - 1,2 - rr,r Enosis misera (Schaus, 1902) F - 12 - rr,r Euphyes derasa (Herrich-Schaeffer, 1869) F,C - 1-3 - rr-rf Eutychide physcella (Hewitson, 1866) F - 4,5 - rr,r Evansiella cordela (Plotz, 1882) F - 12 - rr Hylephyla phylaeus (Drury, 1770) F,C - 2,5 - rr,r Lamponia elegantula (Herrich-Schaeffer, 1869) F,0 - 1,2 - rr lamponia (Hewitson, 1876) F,0 - 2,5 - rr,r

Frequency of Butterflies in Eastern Brazil

37

Lerodea eufala eufala (Edwards, 1869) F,C,0 - 2,3,5,8 - rr-rf

Levina levina (Plotz, 1884) F,0 - 1-3,12 - rr-rf

Libra anatolica (Plotz, 1883) F,0 - 1,2 - r-f

Lucida lucia (Capronnier, 1874) R,F,C,0 - 1,2,4,5,11,12 - rr,r

ranesus (Schaus, 1902) R,F,C,0 - 1-4,12 - rr,r Lycas argentea (Hewitson, 1866) F - 2 - rr

Lychnuchoides ozias ozias (Hewitson, 1878) F,C, - 2,4,5,12 - rr Lychnuchus celsus (Fabricius, 1793) R,F, - 2,5,11,12 - rr-rf Mellana monica (Plotz, 1886) F - 2 - rr Metron chrysogastra (Btl.) (ssp. n.?) F,C,0 - 2 - rr,r

schrottkyi (Giac.) (ssp. tomba Evans, 1955?) F,0 - 2 - rr Miltomiges cinnamomea (Herrich-Schaeffer, 1869) R,F - 1-5,11,12 -

rr,r Moeris remus (Fabricius, 1798) F - 1-3 - rr

striga striga (Geyer, 1832) R,F,C, - 5,12 - rr Naevolus orius orius (Mabille, 1883) R - 5 - r Nastra insignis (Plotz, 1882) R,F, - 1-3,5,12 - r,rf

lurida (Herrich-Schaeffer, 1869) F,C,0 - 2,5,11,12 - rr,r Niconiades caeso (Mabille, 1891) F,C, - 2,5 - rr

merenda (Mabille, 1878) F,C, - 2,5,8,12 - rr Nyctelius nyctelius (Latreille, 1824) R,F,C, - 2-5,12 - rr Orses itea (Swainson, 1831) F - 2,4,5,12 - rr,r Panoquina sylvicola (Herrich-Schaeffer, 1865) F,C, - 2,5 - rr,r Paracarystus evansi Hayward, 1938 C-4-rr Perichares philetes aurina Evans, 1955 R,F,C -4,5-rr,r Phanes aletes (Geyer, 1832) C - 4,12 - rr Pheraeus argynnis (Plotz, 1883) F - 2 - rr

Polites vibex catilina (Plotz, 1886) R,F,C, - 3-5,10,11 - rr-rf Pompeius pompeius (Latreille, 1824) F,C, - 2,3,5 - rr,r Psoralis stacara (Schaus, 1902) R,F,C - 1,2,4,5,10-12 - rr-f Pyrrhopygopsis socrates socrates (Menetries, 1855) F - 11 - rr Quinta cannae (Herrich-Schaeffer, 1869) F - 3 - rr Sabina sabina (Plotz, 1883) F - 5,12 - rr Saliana longirostris (Sepp, 1848) F - 5 - rr Thespieus abatira Zikan, 1938 F,0 - 1-3 - rr-f

ethemides (Burmeister, 1878) F,C,0 - 1,2,8 - rr,r himella (Hewitson, 1868) F,C - 1,2 - rr vividus (Mabille, 1891) F - 2-4 - rr xarippe xarippe (Butler, 1878) F - 1,12 - rr,r Thoon circellatum (Plotz, 1882) F,C - 1,2,5 - rr,r Vehilius clavicula (Plotz, 1884) R,P,C - 1,2,4,5,11,12 - rr,r inca (Scudder, 1872) (?) F - 2 - rr stictomenes (Butler, 1877) R,F,C - 1,3-5,12 - rr-rf Vertica pudor Evans, 1955 F - 5 - rr Vettius artona (Hewitson, 1868) F - 12 - rr

diversus diversus (Herrich-Schaeffer, 1869) R,F,C, -

3-5,8,11,12 - rr,r marcus marcus (Fabricius, 1787) F,C - 2,3 - rr-rf Vidius vidius (Mabille, 1891) F - 12 - rr Vinius letis (Plotz, 1883) R,F,C - 1-5,11,12 - rr,r Virga austrinus (Hayward, 1934) R,F,C - 1-5,11,12 - rr-rf Xeniades chalestra corna Evans, 1955 F,C - 2,3 - rr victoria Evans, 1955 F,C,0 - 1,2 - rr,r

Ebert: Butterflies of Pocos de Caldas

»

Zariaspes mys (Hiibner, 1808) F,C,   -   2,3 - rr

Zenis jebus jebus (Plotz, 1882) F   -   2 - rr

minos (Latreille, 1824) F -   2   - rr 37 Gen. et sp. ign.

inae ( = ,lPyrginae, Section B"; 62 spp.)

Achlyodes mithridates peruvianus Mabille § Boullet, 1.917 (thraso (Jung 1792) is a homonym) F,C,0 - 1,2,4,5,12 - rr,r

busirus rioja Evans, 1953 R,F,C - 2,3,5,10 - rr,r Anisochoria pedaliodes extincta Hayward, 1933 F,C, - 3 - rr,r sublimbata Mabille, 1883 F,C,0 - 2-5,11,12 - rr superior Mabille, 1897 F - 3 - rr Antigonus liborius (Plotz, 1884) (ssp. areta Evans, 1953?) F,C -

1,2 - r Bolla atahuallpai (Lindsey, 1925) C - 2 - rr

catharina Bell, 1937 R,F,C,0 - 3-5,11,12 - rr,r Charidiaempolaeus (Westwood, 1852) F - 5,11 - rr Chiomara asychis (Stoll) autander (Mabille, 1891) F - 3 - rr crenda Evans, 1953 F - 12 - r punctum (Mabille, 1878) 0 - 2,5,12 - r Cogia calchas (Herrich-Schaeffer, 1869) C,0 - 2,3,12 - rr,r

hassan evansi Bell, 1937 F - 5,12 - rr Cycloglypha thrasibulus thrasibulus (Fabricius, 1793) F - 3 - rr Diaeus lacaena lacaena (Hewitson, 1871) F,C - 1-4,12 - rr,r Ebrietas anacreon (Staudinger, 1876) R,F,C - 2,5 - rr Gesta gesta (Herrich-Schaeffer, 1863) F,C,0 - 1-3,5 - rr,r

heteroptera (Plotz, 1884) F,0 - 2,3 - rr Gindanes brebissoni (Latreille, 1822) F,0 - 1,2,12 - rr Gorgopas petale (Mabille, 1888) F,C - 2,5,8,12 - rr Gorgythion begga (Prittwitz, 1868) R,F,C - 1-5,12 - rr-rf beggina beggina Mabille, 1897 F,0 - 1-3,5 - rr escalophoides Hayward, 1941 R - 2 - rr Helias phalaenoides palpalis (Latreille, 1824) R,F,C - 2-5,12 -

rr,r Heliopetes alana (Reakirt, 1868) F,C - 1-3,5,11 - rr,r arsalte (Linne, 1758) F,0-2,3,12 - r laviana (Hew.) libra Evans, 1944 F - 3 - rr omrina (Butler, 1870) R,F,C - 2-5,12 - rr-rf domicella willi (Plotz, 1884) F - 4 - rr Marela thamyroides (Felder, 1867) R - 5 - rr Milanion leucaspis (Mabille, 1878) F,C - 1,2,4,5 - rr MyIon menippus (Fabricius, 1776) R,F, - 2,5 - rr,r Nisoniades bipuncta (Schaus, 1902) R,F,0 - 5 - rr

macarius (Herrich-Schaeffer, 1870) F,C - 3,5 - rr bessus maura Mabille § Boullet, 1916 F - 12 - rr Noctuana diurna (Butler, 1870) F - 5,12 - rr Oechydrus chersis (H.-S.) evelinda (Butler, 1876) F,C - 2,11 -

rr,r rufus Evans, 1953 F,C - 2,11,12 - rr Pachyneuria inops (Mabille, 1877) F - 11 - rr Pellicia dimidiata zama Plotz, 1882 0 - 2 - rr Polyctor polyctor (Prittwitz, 1868) R ^- 4 - rr

Frequency of Butterflies in Eastern Brazil 39

Pyrgus oileus orcus (Stoll, 1780) R,F,C,0 - 1-5,8,11,12 - rr-f

communis orcynoides Giacomelli, 1928 F,C,0 - 2,3,12 - rr,

r Pythonidea lancea (Hewitson, 1868) R,F,C - 1-5,8,12 - rr,r Quadrus cerealis (Stoll, 1782) 0 - 2 - rr

u-lucida (Plotz, 1884) R,F, - 3,4 - rr Sophista aristoteles plinius Plotz, 1882 F - 1 - rr

latifasciata latifasciata (Spitz, 1930) 0 - 12 - rr Sostrata bifasciata bifasciata (Menetries, 1821) R,F, - 1,2,4,5,

12 - rr,r Spathilepia clonius (Cramer, 1775) R - 5 - r Spioniades artemides (Stoll, 1782) R-5-rr Staphylus angulatus Bell, 1947 F - 3 - rr

mazans ascalaphus (Staudinger, 1876) F - 3,5 - rr,r ascalon (Staudinger, 1876) F - 3 - rr chlorocephala (Latreille, 1824) F - 3 - r epicaste epicaste (Mabille, 1878) F - 3 - r incisus (Mabille, 1878) R,F,C, 2,3,5,12 - rr,r Theagenes dichrous (Mabille, 1878) F,C, - 1,2,5,11,12 - rr,r Trina geometrina (Felder, 1867) R,F,C, - 2,3,5,12 - rr-f Viola minor (Hayward, 1933) F,0 - 2,12 - r

violella (Mabille, 1897) F,0 - 2 r Xenophanes tryxus (Stoll, 1780) R,F,C,0 - 2,4,5,12 - r Zera hyacinthinus servius (Plotz, 1884) F - 12 - rr

Pyrrhopyginae (10 spp.)

Croniades machaon (Westwood § Hewitson, 1851) F,C - 2 - rr Granila paseas (Hewitson, 1857) F - 2 - rr Hegesippe luteizona (Mabille, 1877) F - 3 - rr,r Mimoniades versicolor (Latreille, 1823) F - 12 - rr Qxynetra roscius iphimedia Plotz, 1886 C,0 - 2,12 - rr,r Pyrrhopyge charybdis charybdis Westwood § Hewitson, 1852 F,C,0 -

1-3 - rr,r polemon Hopffer, 1874 F - 12 - rr Sarbia antias (Felder, 1859) F,0 - 2 - rr,r

catomelaena Mabille $ Boullet, 1908 F - 2,12 - rr damippe Mabille $ Boullet, 1908 F - 2,11,12 - rr-f

Urbaninae (="Pyrginae, Section A"; 38 spp.)

Aguna albistria albistria (Plotz, 1881) F - 3 - rr asander (Hewitson, 1867) F,C - 2,3 - rr,r megaeles (Mab.) F - 3 - rr williamsi Hayward, 1935 R,F - 4 - rr Astraptes anaphus anaphus (Cramer, 1777) F,C - 1,2,5 - rr elorus (Hewitson, 1867) F - 5 - rr fulgerator (Walch, 1775) F,0 - 2 - rr naxos (Hewitson, 1867) F,C,0 - 1-5,8,11,12 - rr,r creteus siges (Mabille, 1903) F,C, - 1,2,4,5 - rr Autochthon integrifasciatum (Mabille, 1891) F,C - 1,2,5,11 - rr,

r

40                               Ebert: Butterflies of Pocos de Caldas

>

neis (Geyer, 1832) R,F, - 3,5,11 - rr reflexum (Mabille § Boullet, 1912) F - 3 - rr zarex (Hubner, 1832) F,C, - 2,3,5,11 - r,rf Celaenorrhinus punctiger (Burmeister, 1878) F - 3-5,12 - rr-rf

similis Hayward, 1933 R,F,C - 3,4 - rr Chioides catillus catillus (Cramer, 1779) F,C,0 - 2,3,5 - rr,r Epargyreus exadeus exadeus (Cramer, 1779) F - 4 - rr socus socus Hubner, 1825 F,0 - 1,2,12 - rr Phanes vitreus (Stoll, 1781) C - 2 rr Phocides maxima Mabille, 1888 C - 2 - rr

pialia parva Rober, 1925 F - 3 - rr polybius phanias (Burmeister, 1880) F - 2,3,12 - rr Polygonus leo (Gmelin, 1790) F - 2 - rr

manueli Bell § Comstock, 1948 F,C,0 - 2 - rr,r Polythrix caunus (Herrich-Schaeffer, 1869) C - 5 - rr Ridens fulima Evans, 1952 F,C,0 - 5 - rr Typhedanus undulatus (Hewitson, 1867) F - 3 - rr Urbanus doryssus albicuspis (Herrich-Schaeffer, 1869) F - 3 - rr viterboana alva Evans, 1952 F,C - 4,5 - rr dorantes (Stoll, 1790) R,F,C - 2-5,11,12 - rr,r esta Evans, 1952 R,F,C - 2,3,5 - rr-rf evenus (Menetries, 1855) F,0 - 2,3,12 - r,rf evona Evans, 1952 (?) R - 5 - r procne (Plotz, 1881) C,0 - 2,3,5 - rr-rf proteus (Linne*, 1758) F,C - 2-5,12 - rr pronta Evans, 1952 (?) C - 4 - rr simplicius (Stoll, 1790) R,F,C - 2,3,5,12 - rr-rf teleus (Hubner, 1821) R,F,C - 2,4,5,12 - rr,r

Ithomiidae (21 spp.)

Aeria olena Weymer, 1875 R,F,C - 1-5,12 - rr-f Dircenna dero (Hubner, 1823) R,F,C - 1,2,4,5,12 - rr-rf Episcada carcinia Schaus, 1902 R,F,C - 2-5,12 - rr-rf

pascua Schaus, 1902 F,C - 2,5 - rr-rf Epityches eupompe (Geyer, 1832) F,C - 1-5,11,12 - rr,r Hypoleria emyra (Haensch, 1906) R,F - 2,5 - rr oreas Weymer, 1898 F - 1,2 - rr,r salonina (Hewitson, 1855) R - 2,4,12 - r Hypothyris daeta daeta (Boisduval, 1836) R,F - 5,11,12 - rr,r Ithomia agnosia zikani Almeida, 1940 R - 2,4 - rr,r Mechanitis polymnia casabranca Haensch, 1906 R,F,C - 1-5,12 -

rr-ff lysimnia lysimnia (Fabricius, 1793) R,F,C,0 - 1-5 -

rr-f Methona themisto (Hubner, 1818) R,F,0 - 1,2,5,12 - rr,r Placidula euryanassa (Felder, 1860) R,F,C - 2,4,5,12 - rr-rf Prittwitzia hymenaea (Prittwitz, 1865) 1-5,11,12 - rr-rf Pseudoscada adasa (Hewitson, 1854) R,F - 2,4,5,12 - rr-rf

erruca (Hewitson, 1855) R,F - 2,3,5,11,12 - rr-rf Pteronymia carlia Schaus, 1902 - R,F,C - 3-5 - rr,r Thyridia hippodamia (Fabricius, 1775) F,0 - 1,2 - rr

psidii pytho (Felder, 1860) F,0 - 1,2,3,5 - rr,r Tithorea harmonia pseudethra Butler, 1873 F - 2 - rr

Frequency of Butterflies in Eastern Brazil                         41

Libytheidae (1 sp.)

Libytheana carinenta (Cramer, 1777) C - 3 - rr

Lycaenidae (87 spp.)

Plebejinae - (Catochrysopinae - Glaucopsychinae) Hemiargus hanno (Stoll, 1790) R,F,C,0 - 2-5 - rr,r Leptotes cassius cassius (Cramer, 1775) R,F,C - 2,3,5,10,11 - rr (Gen. ign.) cogina (Schaus, 1902) R,F,0 - 2,4,5 - rr

Theclinae ("Thecla" without respect of subgenera)

acaste Prittwitz, 1865 R,C,0 - 2,3,5 - rr,r

agricolor Butler § Druce, 1872 ssp.n. F - 4,5 - rr,r

aphaca Hewitson, 1867 C-2-rr

badeta Hewitson, 1873 C,0 - 2 - rr

bazochii (Godart, 1824) (=thius Hb. ; £=cydia Hew.) F,C - 2-4,11 -

beon (Stoll, 1780) R,F,C - 2,3,12 - rr,r rr,r

bertha Jones, 1912 F - 3 - rr

bolima Schaus, 1902 0 - 12 - rr

bubastus (Stoll, 1780) F - 3 - rr

campa Jones, 1912 F - 2 - rr

cardus Hewitson, 1874 C - 2 - rr

castrena Jones, 1912 0 - 3 - rr

celelata Hewitson, 1874 F,C - 2,5 - rr

celmus (Cramer, 1776) R - 12 - rr,r

hebraea cimelium Gosse, 1880 F - 3-5 - rr,r

conchylium Druce, 1907 F - 3 - rr

cosa Hewitson, 1867 F - 5,12 - rr

crambusa Hewitson, 1874 R - 12 - rr

crolus (Stoll, 1782) F - 2,12 - rr

davara Hewitson, 1868 F,C - 2,11 - rr,r

deniva Hewitson, 1874 F - 1-3,10 - rr

dindymus (Cramer, 1775) R - 12 - rr

ducalis Doubleday § Hewitson, 1852 C - 2 - rr

elika Hewitson, 1867 R - 4 - rr

ellida Hewitson, 1867 R,F, - 1-3,12 - rr

eurytulus (Hiibner, 1819) F,C - 2,12 - rr

faunalia Hewitson, 1874 F,C - 2,4,5,11,12 - rr

geba Hewitson, 1877 F-5-rr

guacanagari Wallengren, 1860 (=azia Hewitson, 1873) F - 2,10,12 -

rr,r hemon (Cramer, 1775) R - 4,5-r

hirsuta Prittwitz, 1865 (=casmilla Hewitson, 1874) F - 3 - rr hygela Hewitson, 1868 R,F - 5,12 - rr,r imma Prittwitz, 1865 (=orcynia Hewitson, 1869 =annia Hewitson,

1874) R,F,C - 3-5 - rr,r janthina Hewitson, 1867 R,F - 12 - r latreillei Hewitson, 1865 F - 5 - rr lisus (Stoll, 1790) C - 2 - rr malina Hewitson, 1867 F,C - 2,4,5 - rr,r

42

Ebert: Butterflies of Pocos de Caldas

marsyas (Linne, 1758) F,C - 3,4 - r

melibaeus (Fabricius, 1793) R,F,C,0 - 1-5,8,11,12 - rr-rf

mulucha Hewitson, 1867 F - 3,12 - rr

(nubilum Druce, 1907) F,0 - 8 - rf not counted)

opisena Druve, 1912 F - 5 - rr

oreala Hewitson, 1868 0 - 2,12 - rr,r

palegon (Stoll, 1780) R,F - 3,5 - rr

(phrosine Druce, 1909 0 - 8 - r not counted)

phrutus (Geyer, 1832) R - 5 - rr

phydela Hewitson, 1867 R,F,C - 1,2,4,5,12 - rr,r

puppius pisidula Druce, 1907 R - 12 - rr

polibetes (Stoll, 1782) F,C - 2-4 - rr

pseudolongula Clench, 1944(?) R,F,C - 1-3,5,12 - rr,r

sangala Hewitson, 1869 F,C,0 - 1-3,5 - rr-rf

schausa Jones, 1912 0 - 4,12 - rr

silumena Hewitson, 1867 F,C - 2,5 - rr

sophocles (Fabricius, 1793) R,F,C - 3-5,8,12 - rr,r

strophius (Godart, 1824) (=panchaea Hewitson, 1869) R,F,C - 3-5,

12 - rr tadita Hewitson, 1877 F,C,0 - 1-5 - rr tarania Hewitson, 1868 F - 3,5 - rr,r tegaea Hewitson, 1868 0 - 2-5,12 - rr-rf tephraeus (Geyer, 1837) R - 12 - rr thordesa Hewitson, 1867 R,F - 5,12 - rr thrasyllus (Geyer, 1837) (thales Fabr. 1793 is a homonym) R,F -

5,12 - rr thyrea Hewitson, 1867 F-5-rr triquetra Hewitson, 1865 F,C,0 - 2-5,12 - rr umbrata (Geyer, 1837) F,C - 5 - rr xeneta Hewitson, 1877 R-5-rr yojoa Reakirt, 1866 F,R,C - 2,5 - rr 20 sp. awaiting determination

Morphidae (5 spp.)

Morpho aega (Hubner, 1822) R,F,C - 4,5,12 - rr,r anaxibia (Esper, 1798) R,F - 2,3 - rr,r epistrophus (Fabricius, 1796) (laertes (Drury, 1782) is

homonym) ssp.n. F,C - 1-3 - rr-rf achilles (Linne, 1758) paulista Fruhstorfer, 1907

R,C - 2,5 - rr,r portis portis (Hubner, 1821) F - 1-3,12 - rr-f

Nymphalidae (76 spp.)

Apaturinae

Doxocopa kallina (Staudinger, 1886) F - 5 - rr selina (Bates, 1865) C - 5 - rr seraphina (Hubner, 1825) R,F,C - 1-5,12 - rr-rf zunilda (Godart, 1821) F - 1,5 - rr

Frequency of Butterflies in Eastern Brazil

Argynninae

Euptoieta hegesia (Cramer, 1779) F - 3 - rr

Biblinae

Biblis hyperia (Cramer, 1779) R,F,C,0 - 2-5 - rr,r Mestra apicalis (Staudinger, 1886) F - 5 - r

Call icon'nae

Callicore sorana (Godart, 1824) R,F,C - 3,5 - rr Diaethria candrena (Godart, 1824) R,F,C - 2-5,11,12 - rr-rf clymena (Cramer, 1775) R,F - 5 - rr eluina (Hewitson, 1854) R,F,C - 2,4,5 - rr-rf Paulogramma pyracmon (Godart, 1824) R - 5 - rr

Catonephelinae

Catonephele sabrina (Hewitson, 1852) F,C, - 1,4,5,12 - rr,r Cybdelis phaesula (Hiibner, 1827-1831) R,F,C - 1,2,4,5 - rr Epiphile hiibneri Hewitson, 1861 R,C - 4,5 - rr,r

orea (Hubner, 1823) R,F,C - 1,2,4,5,12 - rr,r Myscelia orsis (Drury, 1782) R,F - 2,4,5 - rr,r Temenis laothoe meridionalis Ebert, 1965 R - 5 - rr

Charaxinae

Hypna clytemnestra hiibneri Butler, 1866 F - 5 - rr

Memphis arachne victoria (Druce, 1877)

R - 5 - rr appias (Hubner, 1825) R,F,C - 1-5 - rr,r otrere (Hiibner, 1825) R,F,C - 1-5,11,12 - rr,r ryphaea phidile (Geyer, 1837) R,C - 4,5 - rr,r morvus stheno (Prittwitz, 1865) R,F - 1,4 - rr

Prepona chalciope (Hubner, 1823) F - 1,2,11 - rr

Zaretes itys strigosa (Gmelin, 1791) F - 3,5,12 - rr

Coloburinae

Historis odius orion (Fabricius, 1775) R,F, - 3,5 - rr Smyrna blomfildia (Fabricius, 1781) F,C - 2,3 - rr

Eunicinae

Eunica tatila bellaria Fruhstorfer, 1908 F,C - 2,3,12 - rr eburnea Fruhstorfer, 1899 R,F,C - 4,5,12 - rr,r maja (Fabricius, 1775) C - 2 - rr,r margarita (Godart, 1824) R,C - 2-5,12 - rr,r

44                               Ebert: Butterflies of Pocos de Caldas

j

Hamadryadinae

Ectima lirissa lirissa [Godart, 1824) (liria Fabr. 1793 is a

homonym) R,F,0 - 1-5,12 - rr Hamadryas amphinome (Linne, 1767) R,F,C - 3-5,12 - rr,r

arete (Doubleday § Hewitson, 1847) F - 2,5 - rr epinome (Felder, 1867) F,R,C - 1-5,12 - rr februa (Hubner, 1823) R - 3,5 - rr,r feronia (Linne, 1758) (the ?!subsp.'Ms of Fruhstorfer

are synonyms) F - 3 - rr fornax (Hubner, 1823) R,F,0 - 1,2,5,12 - rr

Limenitinae

Adelpha abia (Hewitson, 1850) F - 3,5,12 - rr

calliphane Fruhstorfer, 1915 R,F,C - 1,2,4,5 - rr,r falcipennis Fruhstorfer, 1915 R,F - 1-5,12 - rr,r gavina Fruhstorfer, 1915 R,F - 4,5,11,12 - rr-rf hyas (Boisduval, 1836) R,F,C - 3,5 - rr,r ampla Hayw. mincia Hall, 1938 F,C - 1-5,12 - rr,r mythra (Godart, 1824) R,F,C - 1,2,4,5,11,12 - rr-rf poltius Hall, 1938 R,F,C - 1-5,12 - rr,r isis pseudagrias Fruhstorfer, 1908 R,F - 1,3,4,5 - rr serpa (Boisduval, 1836) F - 2,3,5,12 - rr syma (Godart, 1824) R,F,C - 1-5,11,12 - rr-rf zea (Hewitson, 1850) F - 5 - rr,r Dynamine agacles (Dalman, 1823) R,C - 3-5,,12 - rr,r

artemisia (Fabricius, 1793) R - 5 - rr

tithia (Hubner, 1823) R,C - 2-5,11 - rr,r

meridionalis Rober, 1915 F - 5 - rr

myrrhina (Doubleday, 1849) F - 5 - rr

Marpesiinae

Marpesia chiron (Fabricius, 1775) F - 3 - rr petreus (Cramer, 1776) F - 3 - rr

Melitaeinae

Chlosyne lacinia saundersi (Doubleday § Hewitson, 1847) R,C -

4,5 - rr,f Eresia lansdorffi (Latreille, 1820) R,F,C - 3-5,12 - rr-rf Phyciodes claudina (Eschscholtz, 1821) R,F,C,0 - 1-5,8,11,12 -

rr-f drusilla (Felder, 1861) ssp.n. R,F,C - 2-5,11,12 - rr-f orthia evanescens Rober, 1913 F,C - 1-5,11,12 - rr-rf hermas (Hewitson, 1864) C - 12 - rr ithra Kirby 1910 R,F,C,0 - 1-5,11,12 - rr-rf teletusa teletusa (Godart, 1824) R,F,C - 1-5,8,11,12 -

rr-rf

Frequency of Butterflies in Eastern Brazil 45

Nymphalinae

Anartia jatrophae (Johansson, 1763) F,R,C - 3-5 - rr-rf

amathea roeselia (Eschscholtz, 1821) R,F,C - 1-5,12 -

rr-rf Hypanartia bella (Fabricius, 1793) F,C,0 - 1-5 - rr

lethe (Fabricius, 1793) R,F,C - 1-5 - rr-rf Junonia evarete (Cramer, 1779) R,F,C,0 - 1-5,8,11,12 - rr,r Metamorpha stelenes (Linne, 1758) R,C - 5 - rr Siproeta trayja (Hiibner, 1823) F,C,0-l-5 - rr,r Vanessa virginiensis braziliensis (Moore, 1883) R,F,C,0 - 1-5,12 -

rr,r myrinna (Doubleday, 1849) R,0 - 2,4,5,12 - rr

Papilionidae (16 spp.)

Battus polydamas polydamas (Linne, 1758) R,F,C - 1-4,12 - rr,r polystictes janira (Rothschild § Jordan, 1906) F,C - 1,2,

5,12 - rr,r Eurytides dolicaon deicoon (Felder, 1864) F,0 - 1-3 - rr

lysithous lysithous (Hiibner, 1821) F,C - 1-4,11,12 -

rr-rf Papilio astyalus astyalus Latreille, 1819 F - 3,12 - rr

thoas brasiliensis Rothschild $ Jordan, 1906 F,C - 1,3,

4,12 - rr,r anchisiades capys (Hiibner. 1809) F,0 - 1-3,5,11,12 - rr,r scamander grayi Boisduval, 1836 F,C,0 - 2-5,10,12 - rr,r hectorides Esper, 1794 R,F,C,0 - 1-3,5,11,12 - rr,r torquatus polybius Swainson, 1823 C-2-rf Parides agavus (Drury, 1782) R,F,C - 3-5 - rr,r

bunichus bunichus (Hiibner, 1821) R,F,C - 1-5,10-12 - rr-f nephalion (Latreille, 1819) F,R - 1-3,5,12 - rr-rf proneus (Hiibner, 1827-1831) R,F,C,0 - 11,12 - rr,f Protesilaus bellorophon (Dalman, 1823) F,C - 11,12 - rr,r

sp.ign. (helias (Rothsch. § Jord. 1906)?) F - 3,12 -

rr Pieridae (35 spp.)

Coliadinae

Anteos clorinde (Godart, 1824) F,C - 1,3 - rr menippe (Hiibner, 1816) F,C, - 2,5 - rr Aphrissa statira (Cramer, 1777) F - 2,11 - r Eurema albula (Cramer, 1775) R,F,C,0 - 1,2,4,5,12 rr-rf deva deva (Doubleday, 1847) F,C - 2,5,12 - rr,r elathea (Cramer, 1777) R,F,C,0 - 1-5 - rr-f "jucunda!? (sensu d'Almeida, 1936, nee Boisd. $ Lee. 1833)

lemnia (Felder, 1865) R,F,C - 3-5 - r,rf musa (Fabricius, 1793) R,C,0 - 3-5 - r-f(=gen.hib.of phiale) agave pallida (Chavannes, 1849) F,C - 3,5,7,8,12 rr,r phiale (Cramer, 1775) F,C,0 - 2,4,12 - rr-rf Leucidia elvina (Latreille, 1819) F - 1,2 - rr-r exigua Prittwitz, 1865 F,C - 2,5 - rr-rf pygmaea Prittwitz, 1865 F - 1,2,5 - r

46

Ebert: Butterflies of Pocos de Caldas

Phoebis argante argante (Fabricius, 1775) F - 5 r

neocypris (Hubner, 1823) F,R,C - 2,4,5,11,12 - rr-rf philea philea (Johansson, 1763) F,C - 3-5,11,12 - rr,r sennae sennae (Linne, 1758) R,F,C - 3,5,12 - rr,r Pyrisitia dina leuce (Boisduval, 1836) F - 3 - rr

nise tenella (Boisduval, 1836) R,F,C,0 - 2-5,8,11,12 -

rr-rf Rhabdodryas trite (Linne, 1758) F - 5 - r Sphaenogona arbela arbela (Geyer, 1826) R,F,C - 2,5,8 - rr,r

Dismorphinae

Dismorphia astyocha Hubner, 1827-1831 R,F,C - 1,2,5 - rr,r melite (Linne, 1765) R,F,C,0 - 2,4,5 - rr,r themiesia (Latreille, 1819) F - 2,5 - rr,r

Pseudopieris nehemia (Boisduval, 1836) R,F,C,0 - 1-5,8,11,12 -

rr-rf

Euchloinae

Hesperocharis anguitia (Latreille, 1819) F - 1,2,5 - rr,r

Pierinae

Archonias tereas (Latreille, 1819) F,C, - 2-5,10-12 - rr-ff Ascia monuste orseis (Latreille, 1819) R,F,C - 1-5 - rr-rf Catasticta bithys (Hiibner, 1827-1831) R,F,C,0 - 1,2,4,5,10-12 -

rr-ff Charonias theano (Boisduval, 1836) R - 5 - rr Glutophrissa drusilla (Cramer, 1777) F,C - 1,2,11 - rr,r Melete lycimnia pantoporia (Geyer, 1832) R,F,C - 2,4,5,11,12 -

rr-rf Pereute antodyca (Boisduval, 1836) F - 1-5,11,12 - rr-rf

swainsoni (Gray, 1832) F,C - 1,2,4,5,11,12 - rr-rf

Rtodinidae

Euselasiinae

Euselasia euploea (Hewitson, 1854) R-5-rr eusepus (Hewitson, 1852) F - 5 - rr mys lara Stichel, 1919 R,F,C - 2,5,10,12 - rr hygenius occulta Stichel, 1925 R - 4,5 - rr zara (Hewitson, 1851) R - 12 - rr Riodininae

Anteros formosus (Cramer, 1777) F-2-rr

sp. ign. (n.?) F - 11 - rr Apodemia castanea (Prittwitz, 1865) R,F,C,0 - 2-4,8,11,12 - rr,r

stalachtioides Butler, 1867 F,0 - 2,12 - rr,r Aricoris tutana (Godart, 1822) 0 - 12 - rr Audre epulus signata (Stichel, 1910) 0 - 4,12 - rr,r Baeotis hisbon (Cramer, 1775) C - 2 - rr Barbicornis mona Westwood § Hewitson, 1851 R - 4 - rr Calephelis nilus (Felder, 1861) R,F,C - 3,5,12 - rr,r

Frequency of Butterflies in Eastern Brazil                         47

Calospila bolena (Butler, 1867) F,C - 1,2 - rr

Charis cadytis Hewitson, 1866 R,F - 2-5,10,11,12 - rr-rf

theodora Felder, 1862 R,C - 2,5 - rr,r Chorinea licursis (Fabricius, 1775) R,F - 3-5,12 - rr-rf Cremna alector (Geyer, 1837) F - 12 - rr Echenais sejuncta Stichel, 1910 (?) F - 1,12 - rr,r

sp. ign. F,C - 1,2 - rr,r Emesis diogenia (Prittwitz, 1865) F,C - 2-4,12 - rr fastidiosa Menetries, 1855 F - 3,5 - rr ocypore (Geyer, 1837) R,F,C - 1-3,5 - rr russula Stichel, 1910 F - 5 - rr sp. ign. C - 3 - rr Eurybia misellivestis Stichel, 1910 C - 4 - rr Lasaia agesilas (Latreille, 1805) R,F - 3,5 - rr Lepricornis teras Stichel, 1910 F - 5,12 - rr Melanis jarbas(Fabricius, 1787) R,F,C - 3,5 - rr lycea (Hiibner, 1823) R - 5 - rr marathon (Felder, 1865) R - 5 - rr sp. ign. F - 1 - rr Mesene pyrippe Hewitson, 1874 R,F - 5,12 - rr Mesosemia acuta Hewitson, 1873 (belongs to Eurybia?) F - 2,12 -

rr odice (Godart, 1824) R,F,C - 1-5,10-12 - rr-rf rhodia (Godart, 1824) R,F,C - 1-5,12 - rr,r sp. ign. F - 5 - rr Napaea nepos orpheus (Westwood, 1851) F - 12 - rr Panara thisbe soana Hewitson, 1874 F - 3-5,12 - rr-rf Pterographium satnius (Dalman, 1823) R-5-rr Rhetus periander coerulans Zikan, 1952 F-12-rr Stichelia dukinfieldia(Schaus, 1902) F,C,0 - 2,5 - rr

suavis (Stichel, 1910) F,C,0 - 2 - r Symmachia arion (Felder, 1865) F - 12 - rr Synargis calyce (Felder, 1862)? C - 4 - rr

phillone (Godart, 1824) R,C,0 - 1,2 - rr-rf

Satyridae (34 spp.)

Euptychiinae

Euptychia abretia (Capronnier, 1874) F,C - 2-4 - rr,r ambigua Butler, 1866 C - 2 - r angularis Butler, 1867 C - 2 - r doxes (Godart, 1824) F - 4,12 - r grimon (Godart, 1824) F - 2,11 - rr hemes (Fabricius, 1775) R,F,C - 1-5,8,11,12 - r-ff hygina Butler, 1877 F - 4,5 - r,rf mucosa Butler, 1870 R,F,C - 1-5,10-12 - rr-f necys (Godart, 1824) R,F,C - 1,2,4,5,12 - rr-f ocelloides Schaus, 1902 F - 5 - rr ochracea Butler, 1867 F,C,0 - 2,4,12 - rr-rf pacta Weymer, 1911 0 - 2,3 - rr-rf paeon (Godart, 1824) R,F,C - 1-5,12 - rr,r phares (Godart, 1824) ssp. ign. C - 3,12 - r phronius (Godart, 1S24) R,F,C - 2,4,5,11,12 - rr-f pronophila Butler, 1867 F,C - 2-5,12 - rr-rf

48                               Ebert: Butterflies of Pocos de Caldas

quandtius (Godart, 1821) F,R,C - 1-5,12 - rr-f soter Butler, 1877 F - 3,11 - rr vestigiata Butler, 1867 R - 4,5,12 - rr-rf 10 sp. ign. (all: 0 - 1-4) Taygetis thamyras marginata Staudinger, 1887 R - 4 - r ypthima Hiibner, 1821 F,C - 1-3,12 rr,r

Pronophilinae:

Eteona tisiphone (Boisduval, 1836) F,C - 2,5,12 - rr,r Pedaliodes exul Ihieme, 1905(?) F - 5,12 - rr,r

phanias (Hewitson, 1862) R,F,C - 1,2,4,5,11,12 - rr-rf