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Volume 26, Number 3

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Lugger, O. 1899. Fourth annual report of the entomologist to the state experiment station of the University of Minnesota, to the Governor, for the year 1898. St. Paul, Minn. 280 p.

MAXIMIZING DAILY BUTTERFLY COUNTS

Keith S. Brown, Jr.

Centro de Pesquisas de Produtos Naturais, U.F.R.J., Rio de Janeiro ZC-82, Brazil

Early in 1967, a preliminary draft of a most interesting paper by Heinz Ebert came into my hands for initial comment. This paper (Ebert, 1970) provided me a great impetus toward the systematization of field collecting techniques, and the proper keeping of records. It was especially valuable in pointing out the great rarity of most species in southern Brazil (and probably in most of tropical America), the preponderance of small, inconspicuous, and/or highly localized butterfly species in the Neotropical fauna, and the advantages of having several collectors in an area at the same time to obtain more complete and representative recording of the species present.

I resolved to try to apply the conclusions of Ebert's paper towards a continuing study of the Lepidoptera of the central Brazilian plateau (see Brown & Mielke, 1967, 1968; Mielke, 1967, 1968a, 1968b). Initially, a complete written list was made of the species (including numbers of each sex) that were captured and positively observed1, in each collecting day. The following observation of Ebert (p. 6) provided an initial basis for the effort then made to maximize these daily lists:

"In eastern Brazil the individual frequency of butterflies is generally very low. The success of an excursion is exclusively determined by the number of species found. The higher the number of species found during a trip, the higher the chance to find some regionally (and/or individually) rare species. . . ."

This suggested that the maximization of daily species lists of butterflies, a seemingly unscientific goal (though much employed in a sister area, ornithology), could give a large scientific fallout; and this has indeed proved to be the case. For the benefit of the butterfly-interested public, both amateur and professional, this paper presents a discussion of the methods used for maximization and the results obtained, including a comparison of various Neotropical collecting areas, both in overall Bhopalocera and in individual family or tribal groups.

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Although the collecting methods here described for obtaining maximum daily lists differ in many fundamental aspects from those used by Ebert in his careful long-term studies of the butterfly frequencies in Brazil (which place an emphasis on averages rather than on maximum daily lists), the overall results are very similar and further confirm Ebert?s conclusions on intensive, extensive, and seasonal abundance of the various groups of Brazilian Rhopalocera. This paper thus represents a supplement to Ebert's full discussion, written from a different point of view, and incorporating data derived principally from about 150 collecting days in 1967-1972 (see Table 1 for a selection of these), with maximum expenditure of energy, using methods of intensive or large-scale area coverage, and efficiency and accuracy of recording, developed and sharpened during ten years of taking ornithological censuses.

Methods Equipment

It was ascertained early that the routine use of binoculars during the day's collecting (as in ornithology) not only greatly increased the number of butterfly species that could be positively recorded in a day, but also permitted a beneficial redistribution of energy: most larger common species, which could be unequivocally identified even to the sex at long distance with binoculars, did not need to be captured, and physical effort could be concentrated on collecting the smaller and more unusual species present. In general biological terms, the use of binoculars also allowed careful field studies of high-flying species and unusually wary forms or individuals, rarely mentioned in the lepidopterological literature. In a noteworthy case, the female of a new species of Ithomiinae was observed while laying eggs high in a tree; this observation led to publication of the complete biology of the species together with its original description (Brown & d'Almeida, 1970). Many other immature stages have been discovered in this same manner with binoculars. Also, the nearly extinct Heliconius nattereri was finally relocated, studied, and bred by the author following a chance long-distance sighting with binoculars of a male in Santa Teresa, Espirito Santo (see Brown, 1970, 1972).

An eight-foot pole length with a four-foot extension (for more open areas), easily manageable in all but the densest woods, is necessary to capture the many high-flying species (which generally belong to groups of localized and little-known forms, especially Lycaenidae1 and Hesperi-

1 We use the family Lycaenidae to cover the groxip often discussed as a superfamily, Lycaenoidea (in the Neotropics, the Plebejinae, Theclinae, and Riodininae). Likewise, Nymphalidae — Nym-phaloidea of many authors.

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idae), which can be discovered with binoculars but not always positively identified.

In addition to the binoculars and lightweight net with aluminum pole, minimal collecting equipment is kept (forceps, a belt box with envelopes, a small poison jar in the pocket) to permit rapid movements and efficient handling of captured specimens. Photographic and breeding equipment are left in a central location; early stages discovered are brought back there during "slack" collecting hours in the late afternoon.

Planning

In attaining a maximum list in a given region, or seeking to use a limited number of collecting days most effectively, the lepidopterist should work principally in relatively restricted and disturbed wooded areas (see below), and move around a great deal in the initial explorations. The area chosen must be well reconnoitered for paths and roads, streams and sandy banks, flowers inside and outside the forest, clearings within the woods which catch the sun at various times of the day, and accessible forest edges lit by the first and last sun in the day. The latter areas will provide the majority of the day's lycaenids, and the clearings and flowers the majority of the Hesperiidae. A partly cleared hilltop will provide dozens of additional species throughout the day in both these groups, plus many additional and unusual Papilios and Nymphalidae. Baited traps (as many as 30) should be placed along trails and on woods-edges on the evening or night previous to a collecting day. By using traps to capture the many bait-attracted nymphalines and satyrines (a number of which enter during the twilight and dawn hours), the collector can focus his attention on the smaller species which do not catch themselves so easily.

Collecting Period and Distribution of Energy

In the ideal case, weather and his strength permitting, the collector should work from sunup to dark, leaving the woods edge at sundown (when the Lycaenidae finally sit down for the night) to capture Bras-solini at their suspected or prelocated fly ways. Thus the condition of maximum effort may be fulfilled, giving the over-all results a measure of consistency. "Slack periods" in the early morning and late afternoon, or during cloudy spells, may be used for resting, moving between collecting areas, and harvesting the traps. In very hot lowland tropical areas (the coastal plain in southeast Brazil in summer, much of the Amazon Basin near the rivers, or lowland Panama) both butterflies and collectors are often driven into inactivity by the heat between 1100 and 1430; but in

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hilly or mountainous regions, both can remain active during most of the daylight hours, with peaks in late morning and mid-afternoon.

A single collector, for best maximization, should move rapidly from habitat to habitat (using a vehicle such as a Jeep to cover greater distances), collecting a number of times during the day in each subarea of the locality chosen. On the other hand, a party of collectors will get more results and tire less during the day if they divide up the locality, each covering one biotope or contiguous area very thoroughly during the entire day.

In view of the great preponderance of small-sized Hesperiidae and Lycaenidae in the Brazilian butterfly fauna (see Table 1, end), the effort to maximize lists requires a heavy concentration on these often ignored groups, using binoculars to locate high-perching individuals and spending much time collecting the many species that visit flowers. As Ebert (1970) has noted, the vast majority of the species in these two families have a very low intensive frequency; even in areas where over 300 species in each family have been recorded over long periods, the maximum day's list under optimal conditions rarely passes 100 species in either family. Casual (non-maximized) collecting usually yields but 10-40 species of Lycaenidae or Hesperiidae in a day; concentration on these groups can raise the levels to 30-80 or more (see Table 1).

Results

The comparison of results obtained in completing the list for the central plateau before and after the adoption of these methods is instructive. In six weeks' collecting in the planalto, before the use of the maximization method, the author and Nirton Tangerini added only 25 species to the total published list (Brown & Mielke, 1967). Using the methods to maximize daily lists, and thereby collecting preferentially the less noticeable species, the author with Stan Nicolay or Karl Ebert added nearly 300 species to the list in another six weeks' collecting in 1967-1970. The total list for the planalto is now over 950 species (see Table 1); most of the added species are small or inconspicuous forms (Satyrinae, Lycaenidae, and Hesperiidae) which probably would have been passed over in casual collecting.

In two weeks' collecting in Belo Horizonte using these methods in May 1967 (the first real test of maximization), the author added 145 species to the published list for the area (Brown & Mielke, 1967). On one sunny day, 260 species of Rhopalocera were positively identified, over 90% of these being caught.

On the local scale, the maximization method could be used to confirm

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a fact well-known to Neotropical lepidopterists but often misunderstood in other areas: the most rich and, surprisingly, often most representative collections are to be made in highly restricted areas of partly cut forest, surrounded by fields and swamps, preferably with many low flowers, and liberally served by paths, roads, and clearings. While the legendary (and often imaginary!) areas of vast virgin tropical forest invariably harbor many unusual and characteristic species, these tend to occur in very low density throughout the forest, and must seek sun at levels unattainable by the collector's net. The best overall collecting is always in small varied woodlots where most species thrive and remain accessible.

On the regional scale (see Table 1), the maximization method revealed that the largest intensive species frequencies for daily lists (though not necessarily for long-term censuses), could be found in the blend zones between the cerrado area of the Brazilian planalto and the adjacent tropical forests of the southeast coastal mountains (as; in Belo Horizonte or Pocos de Caldas) or the Amazon Basin to the north. The rarer species seemed to be more common in these areas, probably because of the highly varied microclimates and floral niches present in the hilly transition areas between different faunal regions.

In the northwestern blend area between the cerrado and the upper middle Amazon region in central Mato Grosso, preliminary application of these methods in the summer of 1969 suggested that intensive winter collecting might give lists approaching the limits of daily butterfly numbers for the Neotropics. Therefore, when in May 1969 an excursion to the Chapada de Guimaraes was made in company with Stan Nicolay, full plans were developed for intensive collecting with maximization methods, both along the way and in the Mato Grosso blend area. The results of this trip (see Table 1) amply confirmed the exceptional position of the Chapada de Guimaraes in central Mato Grosso with respect to relative butterfly abundance. This area produced in one day a list of 361 species (307 recorded by the author alone), which, for the time being, stands far above the daily lists obtained by us or others in any other area of the Neotropics or, indeed, in the rest of the World.2

2 The late Dr. Richard M. Fox claimed to have collected "about 350" species in a single day's collecting near his field station in Liberia (personal communication). As he reports only 450 species for the whole country (Fox, Lindsay, Clench & Miller, 1965), we must conclude that, unless the percentage of rare and localized species in the Liberian fauna is vastly lower than that in the Brazilian fauna, this estimate may be somewhat exaggerated. The author also has a letter from John H. Masters indicating that a list of 350 would be attainable in a single day's collecting in the Amazon Basin. We presume that this would be in the blend areas on its western edges, where the highland and basin faunas meet and mix. The author has visited a number of such areas in Colombia, Peru, Ecuador, and Bolivia; several of them offered the same subjective impression of extreme butterfly abundance that is evident in central Mato Grosso.

A day's total of 350 should be near the limit for a single collector even in such exceptionally rich areas, not due so much to butterfly abundance as to limitations on the speed of moving between ecological niches, and observing, collecting, and enveloping specimens. In some exceptionally varied blend areas on the borders of the Amazon Basin, a party of three or four collectors might be able to pass the limit of 500 species in a single day.

Locality

Table 1. (Continued)

Date

Weather

No.

Flowers col. mo be. sat da ith

h++

++

5 18

TOTALS for mountain forests near Rio, 28 days

Generally depauperate daily fauna, though f ariety in the smaller species. Peak in m

8 10 32 4 26

Santa Teresa, Espirito Santo: Heavy, steep virgin forest, cold in winter, hot in summer.

luiAU ror santa JLeresa area, 1UU collecting days

Conceicao da Barra, Espirito Santo: Heavy dense virgin forest.

Pedro Canario, Espirito Santo: Remnant of same forest, with flowering fields.

Mirassol, Sao Paulo: Semitropical deciduous forest.

23-VI-68        Clear, warm

4-V-69         Clear, hot

27-VII-67 Clear, warm 20-VI-68        Clear, hot

5-IX-

Clear, hot

Table 1. (Continued)

Table 1. (Continued)

Locality

Date

Weather

No.

Flowers col. mo br sat da ith a

Part cloudy, warm

Jardim Zoologico,                 24-II-6

D.F.: Low humid forest, marsh,

swamp.                                                            NOTE:

TOTALS for Brasilia area, 43 collecting days

++

15

Peak in autumn. Abundance of Satyrinae 4 13 45 3 22

REGIONAL TOTALS over many years' collecting Eastern Pernambuco—Lowland tropical forest (HE) Eastern Sao Paulo—Lowland and mountain forest (E) Rio de Janeiro and vicinity (low & mt. forest) (HE) Rio de Janeiro and vicinity (author's totals) Cerrado portion of Planalto (excl. blend zones) Median-littoral humid forest (Bahia, Esp. Santo) TOTALS FOR EXTRA-AMAZONIAN BRAZIL

(First three

7 23 3 15

7 19 46 5 32

19     39

20     43

17 75 22 70 37 150*

26 38 29 46

75

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Regional and Seasonal Comparisons by Subgroup (Table 1)

It is of interest to note here the seasons and areas in Brazil, and in other regions in the Neotropics, where, in our experience and based upon these methods of intensive collecting, there may be expected maximum numbers of species in each of the major groups of diurnal Lepidoptera.

Morphos in southern Brazil are most common in Santa Teresa, Espirito Santo; all eight species which we recognize in the extra-Amazonian area may be found flying together there in April. Very few other such small areas harbor over six species, and in more subtropical climates these rarely overlap during their single yearly broods in summer. If the same criteria of conspecificity are applied in other regions, eight species may also be captured (though possibly not all at the same season) in some areas of the eastern slopes of the Andes in Peru; and seven species may be found flying together at one time in many warmer parts of tropical America, where most species are multibrooded.

Satyrines (including brassolines) seem to be most diversified in areas of alternating dry scrubby woodland and moist riparian forest, with a strong additional element of open grassland and marsh, such as the Chapada de Guimaraes and Brasilia areas, with 54 and 58 species known to date. Some larger areas have the required diversity to harbor nearly 100 species in this subfamily. A few areas in the Andes may surpass 100 species, though it seems unlikely that more than fifty could be captured in a day. The brassolines alone seem most common in hilly areas where neighboring faunal regions meet; the key to their observation and collection is always the discovery of their preferred afternoon and evening fly ways and the liberal use of traps with preferred local bait. The area of Santa Teresa has at least twenty species, of which ten have been observed in a single day; we know of no other area in the Neotropics with a similar species abundance. While satyrines and brassolines in general show less strict seasonal variation than other nymphalid groups, individual species often show marked alternations of heavy broods and essentially complete absences. A few temperate-zone species are single-or double-brooded, in summer only.

In Brazil, the maximum number of species of Ithomiinae is reached in the median-littoral region, a broad area of moist coastal tablelands in southern Bahia and northern Espirito Santo which shelters many endemic species. Here, 30 species can be found in a single "pocket" in winter, and 25 to 27 can reasonably be collected in a single day. These numbers do not compare with those found in the upper Amazonian and Orinocan tributaries on the slopes of the Andes in Bolivia, Peru, Ecuador, Colombia, and Venezuela, clearly the principal center of geohistorical evolution in

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the subfamily. In the Chanchamayo valley (La Merced), Peru, and in north-central Venezuela, the author discovered nearly 50 species present, of which 40 could reasonably be captured in a single day, especially with the use of dried Heliotropium plant (Beebe, 1955; Masters, 1968) hung at favored locations within the "pockets." Collection of ithomiines is generally best in the dry season, though a few montane species are most common in the warmer, moist summer season. Many species in the family are notable for their accentuated and unpredictable variations in populations and abundance from season to season and year to year. The localized dry season "pockets," in which all of the species tend to fly together, are much diluted during the wet seasons, and occasionally switch their location dramatically from one year to the next.

The maximum number of species of heliconians found in one locality of extra-Amazonian Brazil is 17, in Santa Teresa; 16 of these have been observed in a single day. Again, this compares poorly with the numbers present in some parts of the Andean slopes of Ecuador and Colombia; on the Rio Negro in Meta, Colombia, over 30 species have been found, of which 22 were observed by the author in two days (Brown & Mielke, 1972). Nymphalines, many of them attracted to bait, have been found most commonly in extensive and very moist, slightly disturbed forests at the base of the mountains near Rio, and in the interior near Brasilia, Buriti (Chapada de Guimaraes), and Rio Vermelho (Amazon-type highland forest in western Mato Grosso), but the total lists for an area do not vary much from an average of 80 (locality) or 110 (larger region). The greatest abundance of Nymphalidae, broadly speaking, has been observed in Rio Vermelho (161 species in one day); however, if total numbers observed (as a smaller number per day) over a long period is considered, Santa Teresa, which blends four major faunal regions, is much richer.

Almost all tropical nymphalines, including heliconians and charaxines, are to be found most commonly in late summer and fall in cooler areas, and late winter in hotter regions. The genus Eunica contains many species that are practically limited to one brood in the late winter. A few larger species of Charaxinae are double- or triple-brooded only, showing much variation in abundance and freshness of individuals caught at different seasons.

For the extensive collection of Theclinae, no area can compare with the Brazilian cerrado in the early dry season. Although this habitat has much akin faunistically to an open forest, the trees rarely exceed 20 feet in height, and thus treetop-loving species can be netted easily, together with those preferring low flowers and grass. The spacing of the low trees assures abundant sun all day long, and the interspersal of occasional

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denser growth (cerradao), more open areas (campo cerrado), and moist riverside and headwater woods guarantees the presence of a very wide variety of Theclas. The 47 species in this subfamily captured by the author and Karl Ebert in Itirapina, Sao Paulo, a southern outpost of the cerrado within the blend zone, in May 1967 (Ebert, 1970) represented an early and successful test of some of the methods described in this paper. In a good year (such as was 1972) in the Brasilia area, a day's list of nearly 100 Theclas should be attainable by a party of three or four collectors; the maximum to date, with two persons, is 84, and a total lycaenid list of 137 species.

The only areas we have seen to match the forests of the Chapada de Guimaraes for Riodininae are the lowland woods near Manaus, where 50 species were captured in two days in January 1970, and the swampy woods near Belem, where 45 species were found in two days in late December, 1970. Presumably, an area could be found in the lower middle Amazon which would produce nearly a hundred species of these brightly colored small butterflies in a day. Diversity of flora is a good clue to abundance of Riodininae, and favored male patrolways (hilltops and open moist woods are best) must be located, and hours of flight activity ascertained, to give good collecting.

In Brazil, Papilionidae are most abundant at the very beginning of the rainy season in September, especially in the upper Paraguay River basin, where over 20 species could be captured in a day; similar or greater numbers should be attainable in many areas of the upper Amazon basin, as well as in the foothills near Rio de Janeiro at this same season (24 species recorded in Xerem, not all on one day). The fall and winter are very poor in Papilionidae, except for occasional Parides in warmer areas.

Pieridae have been found most abundantly in the same areas and seasons as Papilionidae; the diversity is greatest in the foothills near Rio where mountain and lowland species mix (44 species recorded, 22 in a single day in Xerem). Outside Brazil, somewhat greater numbers may be encountered in favored seasons on the eastern slopes of the Andes,

Hesperiidae appear commonly in almost all Neotropical areas with diversified ecological biotopes (though much more rarely in extensive virgin forests); a daily list of over 100 species should be attainable in any of a number of regions in the good flower seasons. Table 1 shows an extraordinary correlation of numbers of skippers captured with flower abundance, which should be taken to heart by those who specialize in this family. Prominent in the flower groups that bring these butterflies out of the forest and concentrate them in accessible areas are members of the Eupatoriae, which flower principally in fall and winter in Brazil.

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Summary

A method for the conscious maximizing of selected daily butterfly lists in southern Brazil has permitted much new information to be obtained on the occurrence and biology of the small, localized, and/or high-flying species in the fauna, which tend to be least known. It also has provided a reasonably standardized basis for comparisons between different localities and seasons, both in overall number of species present and in relative abundance of the various subgroups of Rhopalocera. General conclusions on extensive, intensive, and seasonal frequencies closely match those presented by Ebert (1970), though the collecting methods used are fundamentally different.

Acknowledgments

The author is indebted to Dr. Heinz Ebert for the original stimulation of the work leading to this paper, as well as its revision at several stages of development. The collecting days described in Table 1 were realized with the assistance of a number of experienced and uncomplaining colleagues, especially Dr. Ebert and his son Karl, Olaf Mielke, Claudionor Elias, Stan Nicolay, Nirton Tangerini, and Luis Otero; many other persons aided in the localization and exploring of the areas mentioned. Outside Brazil, the author was priveleged to be introduced to the Andean fauna by Dr. and Mrs. L. W. Harris (Peru), Dr. E. W. Schmidt-Mumm (Colombia), and Drs. K. Negishi and S. S. Tillett (Venezuela). Financial support of the chemical study of Brazilian insects by the Conselho Nacional de Pesquisas (of which the author is honorary lecturing investigator, ''Pesquisador-Conferencista"), the Banco Nacional de Desen-volvimento Economico, the Conselho de Pesquisas e Ensino para Gradua-dos of the U.F.R.J., and the National Science Foundation (Grants GB 5389 X and GB 5389 XI), is gratefully acknowledged.

Literature Cited

Beebe, W. 1955. Two little-known selective insect attractants. Zoologica (New

York) 40: 27-32. Brown, K. S., Jr. 1970. The Heliconians of Brazil (Lepidoptera:Nymphalidae).

Part I. Rediscovery of Heliconius nattereri in eastern Brazil. Entomol. News. 81:

129-140. ----------. 1972. The Heliconians of Brazil (Lepidoptera :Nymphalidae). Part III.

Ecology and biology of Heliconius nattereri, a key primitive species near to

extinction, and comments on the evolutionary development of the species of

Heliconius and Eueides. Zoologica (New York), in press. Brown, K. S., Jr. & O. H. H. Mielke. 1967. Lepidoptera of the Central Brazil

Plateau. I. Preliminary list of Rhopalocera. J. Lepid. Soc. 21: 77-106, 145-168. ----------. 1968. Lepidoptera of the Central Brazil Plateau. III. Partial list for the

Belo Horizonte area, showing the character of the southeastern "blend zone."

J. Lepid. Soc. 22: 147-157.

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----------. 1972. The Heliconians of Brazil (Lepidoptera: Nymphalidae). Part II.

Introduction and general comments, with a supplementary revision of the tribe.

Zoologica (New York) in press. Brown, K. S., Jr. & R. Ferreira D' Almeida. 1970. The Ithomiinae of Brazil

(Lepidoptera: Nymphalidae). II. A new genus and species of Ithomiinae,

with comments on the tribe Dircennini d'Almeida. Trans. Amer. Entomol. Soc.

96: 1-17. Ebert, H. 1970. On the frequency of butterflies in eastern Brazil, with a list of

the butterfly fauna of Pogos de Caldas, Minas Gerais. J. Lepid. Soc. 23: supplement 3, 1-47. Fox, R. M., A. W. Lindsay, H. K. Clench & L. D. Miller. 1965. The Butterflies

of Liberia. Mem. Amer. Entomol. Soc. 19: 1-438. Masters, J. H. 1968. Collecting Ithomiidae with Heliotrope. J. Lepid. Soc. 22:

108-110. Mielke, O. H. H. 1967. Lepidopteros do Planalto Central Brasileiro, IV. Notas

complementares sobre os Hesperiidae da regiao e descrigao de cinco especies

novas (Lepidoptera, Hesperiidae). Rev. Brasil. Biol. 27: 125-134. ----------. 1968a. Lepidoptera of the Central Brazil Plateau. II. New genera, species

and subspecies of Hesperiidae (Rhopalocera). J. Lepid. Soc. 22: 1-20. ----------. 1968b. Lepidoptera do Planalto Central Brasileiro. V. Novas especies de

Hesperiidae e anotacoes sobre outras especies conhecidas. Rev. Brasil. Biol.

28: 447-455.

AN UNUSUAL MOTH IN CENTRAL ILLINOIS

After working on my car, at about 1400 on 14 November 1971, I drove it through a parking lot across the street from my home. I then noticed a large moth clinging to a building, about ten feet off the ground. A closer look revealed that it was a large noctuid. Immediately, I opened the back of the car, grabbed my net, made a lunge at the moth, and managed to capture it.

As soon as I got home, I checked my revised edition of Holland's Moth Book, and found that the moth I had just caught was Thysania zenobia Cramer. After reading that it was a "South American moth," I knew that I had caught something quite unusual for central Illinois. However, a moderately strong wind had been blowing from the south for several days preceding the capture, and as the specimen was slightly worn, this probably explains its presence this far north, especially at such a late time of year.

I am sure that this capture does not constitute a state record after checking with the University of Illinois and Eastern Illinois University; but it might be a new record at least for Coles County. If any members know of a previous capture of this species in this county, I would sincerely appreciate hearing from them.

Charles K. Swank, 616 S. 15th St., Mattoon, Illinois 61938.

WING-SHAPE AND ADULT RESOURCES IN LYCAENIDS

Various small species of blues (Lycaenidae: Lycaeninae) are characterized by rather narrow elongated wings—a Nearctic example being Zizula cyna Edwards. In September of 1971 some observations were made on a related African species, Zizula hylax (F.) in the Gombe Reserve on Lake Tanganyika, Tanzania.