STUDIES ON THE FAUNA OF CURAÇAO AND OTHER

CARIBBEAN ISLANDS: No. 185

The caddis-flies (Trichoptera) of

Cuba and of Isla de Pinos: a synthesis

by

L. Botosaneanu

(Instituut voor Taxonomische Zoologie, Universiteit vanAmsterdam)

the fauna of Even a few years ago, caddis-fly remained very

poorly known. The situation notably improved, especially following

the of collections made the either study throughout Island, by my-

self, or by Cuban entomologists (Botosaneanu & Sykora 1973,

Botosaneanu 1977, Botosaneanuin press).

somewhat less and At present, than 80 species subspecies are known from Cuba (2 Rhyacophilidae Hydrobiosinae, 8 Glossosoma-

tidae Protoptilinae, 6 Philopotamidae, 1 Psychomyiidae Xypho-

centroninae, 6 Polycentropodidae, 6 Hydropsychidae Hydropsy-

chinae and 3 Macronematinae, 31 Hydroptilidae, 3 Leptoceridae, 2

Odontoceridae, 4 Calamoceratidae, 5 Helicopsychidae). But we can

expect some 10 additional species to be discovered. These figures

when those obtained for the are reasonably high compared to other antillean Islands (see especially Flint 1964, 1968a, 1968b): for

Jamaica and , both well investigated and smaller than

39 and Cuba, the figures are 32 respectively; only 27 species were

quoted from (, certainly having a rich fauna, is still very poorly known); 44 species have been reported on the whole for the Lesser Antilles. The percentage of endemical taxa is remark-

ably high: 61 from the 76 total. These are species of Atopsyche

Bks., Cubanoptila Sykora (genus endemical in Cuba), Cariboptila

Flint, Campsiophora Flint (2 purely Antillean genera), Polycentro- pus Curt., Hydropsyche Pict., Smicridea McL., Leptonema Guérin, 34

Macronema Pict., Alisotrichia Flint, Ochrotrichia Mos., Metrichia

Ross, Loxotrichia Mos., Oxyethira Eat., Hydroptila Dalman, Neotri-

chia Mort., Oecetis McL., Marilia F. Müller, Phylloicus F. Müller,

Helicopsyche Siebold.

CASES OF VICARIANCE (see Table)

Cuba - like the Antilles the whole — land of represents on a pro-

mise for of the results of isolation1 a study geographic ); investiga-

tions carried on its caddis-fly fauna allow to distinguish a lot of

of cases vicariance (at the generic, specific, or subspecific level): i.

between the Greater Antilles considered as a whole, and other areas of the ; 2. between different islands of the Greater An-

different natural of within tilles; 3. between regions Cuba; 4. or even

the limits of one of these natural regions. Selected are here a few

examples offered by the Cuban fauna for illustrating these 4 dif-

ferent Occ. cases (abbreviations: C. = Cuba; = Occidental region;

Oriental = Central I.P. = Isla de Or. = region; Centr. region;

Pinos; see Fig. 10).

- Bots. - Ross 1. Chimarra guapa (C.) C. patosa (); Loxotri-

chia glasa Ross (I.P., southern U.S.A.) - L. dalmeria Mos. (Mexico);

Oxyethira alaluz Bots. (C.) - O. maya Denn. (Georgia, Florida);

Hydroptila medinai Flint (C., Puerto Rico) - H. mexicana Mos.

(Mexico); Neotrichia iridescens Flint (Greater and Lesser Antilles) -

N. olorina (Mos.) (Mexico).

When compared to the trichopteran fauna of the neighbouring

of Cuba zones Continental America, that of is clearly distinct through

but some positive features, especially through some negative ones, being (like that of the other Antilleans) a notably impoverished fauna. The low Hydrobiosinae are represented only by a very num-

The ber of species belonging to only one genus, Atopsyche. Protop-

rather of 3 tilinae form a good bunch species, belonging to only

that "... the inter-American and has been 1) Ross (1967) says circum-Caribbean area

the most hotbeds of evolution and and that "The one of World's prolific speciation",

of the inter-American can be considered a geotectonic oscillations area ... species- making machine of unusual efficiency". 35

all of endemicalfor the Greater Antilles. The Chimarra genera, them

few but rather are relatively phyletically varied. The Polycentropo-

didae are rather well represented (2 Antillopsyche and 3 Polycentro-

pus).

One of the remarkable the most positive features is presence of a

group of 4 nearly related Hydropsyche (these are - together with 2

species from Hispaniola - the only representatives of the genus in

the the otherwise absent from South Antilles, genus being America).

Excepting genus Alisotrichia, present with 5 species at least, the

“ of the Leucotrichia-group” well else- hydroptilids , represented

the where, are practically lacking. The list of remaining hydropti-

lids is reasonably long and "normal", but the Hydroptila, so much

diversified for instance in Mexico, are represented only by a small

of the it is number species belonging to great “consimilis-group” 1 ;

between the hydroptilids, too, that some of the most phyletically

isolated Cuban species can be found: Ochrotrichia insularis Mos.,

O. islenia Bots., or the 3 species of Neotrichia (alata Flint, pequenita

Bots., pinarenia Bots.). There are neither Brachysetodes nor Tri-

plectidinae among the Leptoceridae. No special comment is neces-

for Odontoceridae for the Calamoceratidae. sary the or A certain

"lustre" is confered to the Cuban fauna by the Helicopsyche:

5 1 2 of them - H. at least species (but probably or more), one gran-

piedrana Bots. & Syk. - being particularly isolated.

2. - Atopsyche vinai Syk. & Bots. (C.) — A. taina Flint (Hispani-

ola); Chimarra cubanorum Bots. (C.) - C. spinulifera Flint (Hispa-

Bks. and A. - niola); Antillopsyche wrighti (C.) aycara Bots. (C.)

A. tubicola Flint (Puerto Rico); Alisotrichia fundorai (Bots. & Syk.)

(C.) - A. hirudopsis Flint (Puerto Rico) and perhaps A. argentilinea

Flint Ochrotrichia insularis Bots. - (); ayaya (C.) O. insularis

insularis Mos. selvatica Bots. - H. (Jamaica); Hydroptila (C.) an-

cystrion Flint (Jamaica). Several cases of incipient subspeciation ha-

been discovered ve (Hydroptila, Oxyethira).

The affinities between the Cuban fauna and that of each of the other islands of the Greater Antilles, are approximatively of the

same order. 36

- divided Cuba J. The cuban geographer NUNEZ-JIMENEZ (1972)

in 5 "natural regions": Occidental, Central, Camagiiey-Maniabon,

Oriental, and Isla de Pinos. This division is certainly warranted,

the fauna. the as shown also by With the exception of

known Camagiiey-Maniabon region (almost no caddis, and certainly

each a fauna a number very poor fauna), region possesses including of characteristic elements. The richest and best characterized region

known is the Oriental one: 32-33 taxa exclusively from this region.

The 8-9 5 Occidental region follows at a good distance, with taxa.

Isla taxa are exclusively known from the Central region, and 2 from

de Pinos. All this very well reflects the geomorphological, geologi-

cal, and hydrological peculiarities of the different regions.

Rather frequently, a species inhabits Occ. + Centr. (4 cases), Occ.

I.P. 4 -)- (3 cases), or these 3 regions together (2 cases); there are

species inhabiting Or. -f- Centr.; several species are widely distri-

buted the and the be consi- throughout Island, following ones may

dered as being the most frequent and widely distributed: Chimarra

Bks. (Curgia) pulchra (Hag.), Smicridea comma (with some doubt!

further see below), Neotrichia iridescens Flint, Nectopsyche cubana

(Bks.), Oecetis inconspicua (Walk.). To come up again to the vica- riants, this time between different natural regions, the following

examples will be quoted: Cariboptila soltera Bots. (Occ.) - C. guaji-

Bots. and Bots. Macronema Bks. ra C. poquita (Or.); gundlachi

(Occ. -(- Centr.) - M. tremenda Bots. (Or.); Alisotrichia flintiana

Bots. (Or.) - A. cimarrona Bots. (Occ.); Ochrotrichia caramba Bots.

(Occ., Or.) - O. villarenia Bots. (Centr.); Metrichia espera Bots.

(Occ.) - M. cafetalera Bots. (Centr.) - M. munieca Bots. (Or.); Phyl-

loicus chalybeus chalybeus (Hag.) (Or.) - P. chalybeus ssp. (Occ. +

I.P.); Helicopsyche comosa Kings. (Occ.) - Helicopsyche n. sp. prope

- be lutea comosa may Hag. (Or.).

- A remarkable of that 4. case non-geographic vicariance is of

two related Alisotrichia and very nearly species: chiquitica Bots.,

A. alayoana Bots. Both of them are inhabitants of the Oriental re-

in gion but they apparently never occur together the same water

course (competitive exclusion?). 37

TAXONOMICAL AND OTHER PROBLEMS

I have provisionnally reported most of theSmicridea taken in Cuba

but I forced that to the species comma Bks.; am to say practically

each locality, or group of localities, has given a slightly different

morphological type, so that the specific determination remains

doubtful, especially in the absence of dry specimens; Dr. O. S. FLINT,

find the solution of this the other jr., proposes to puzzle. On side, a

quite similar problem is that of a lot of Helicopsyche "populations"

in should be attached the “haitiensis- which my opinion to group”

Prof. H. H. Ross who is Cuban Heli- (according to studying my

copsyche: "They are definitely the most puzzling collections of

Helicopsyche I have ever studied").

The in rather Cuban of androco- occurrence many Trichoptera nial formations and of specialized hairs should also be emphasized.

For instance, Cubanoptila muybonita Bots. is, as far as I know, the

only described Protoptilinae to have in the fore wings of the <$ a

wide zone covered with small, highly specialized hairs; Ochrotri-

chia caramba Bots., and O. villarenia Bots. are probably the only

in - representatives of the genus to have scales the fore and hind

wings of the Oecetis maspeluda Bots. is readily distinguished by

the of fine black hairs in the well groups hind wings (<£, $) as as by

the occurrence ($) of a very long penicillus of black androconial

hairs inserted at the base of the anterior femur; the Helicop-

of the taken in least in the syche “haitiensis-group” at one locality

Oriental region, have on their anterior wings a round androconial

area covered with black scales; Helicopsyche comosa Kings, and a

in the of new, not yet described, species, are certainly unique limits

the the dense hairs their genus, by covering of long on gonopods.

GEOGRAPHICAL DISTRIBUTION (SEE TABLE)

the The close analysis of geographical distribution on one side, of the "sister relationships" on the other side, leads to the following conclusions (see also Ross 1967 and FLINT 1977).

— number of bear a) A great species a quite distinct "Antillean

which stamp", belonging to genera or to species groups are strictly 38

all Antillean (or even Cuban). Some examples: the Protoptilinae,

Chimarra (C.) cubanorum Bots. and C. (C.) garciai Bots., the two

Antillopsyche, Polycentropus nigriceps Bks. and P. turquino Bots,

Alisotrichia fundorai (Bots. & Syk.), A. chiquitica Bots. and A.

Ochrotrichia insularis and O. islenia alayoana Bots., ayaya Bots.

Bots., Helicopsyche granpiedrana Bots. & Syk.

b) -If the somewhat more remote affinities of the taxa are also considered, we find that, in many instances, the taxa show more or less distinct relationships with elements of continental Central Ame- rica and/or of S (especially SE) North America (if there is not simply identity). Some examples: Polycentropus criollo Bots., all the

Hydropsyche, Leucotrichia cf. tubifex Flint, Ochrotrichia caramba

Bots. and O. villarenia Bots., Orthotrichia americana Bks. and O. cristata Mort. Loxotrichia glasa Ross, the 5 Oxyethira, the 4 Hydrop- tila, all the Helicopsyche excepting granpiedrana.

- On the the c) contrary, taxa certainly or possibly belonging to trends of South American origin, are extremely seldom, but never- theless highly interesting. Examples: Atopsyche cubana Flint and

A. vinai Syk. & Bots., Chimarra (Curgia) pulchra (Hag.) and C.

(Curgia) alayoi Bots., C. (C.) guapa Bots. It is now a demonstrated fact that the trichopteran fauna of Cuba does not belong to the

"faunal circle" represented in the Lesser Antilles northern

South America + southern continentalCentral America, but to that including the Greater Antilles + southern North America + north- ern continental Central America.

Much interest is being now displayed for the Mexico Bay-Carib-

bean Sea much and areas, including geological, geomorphological, paleogeographical research, the results of these studies having im- plications on the problem of the origin and evolution of the fauna inhabiting the lands of the area. More and more publications are accumulating on this item (see, for instance, the comprehensive

of ROSEN with conclusions often di- paper 1975), being extremely

the taken into the vergent depending on group account, on more or less sound character of the study, on the geological and paleogeo- graphical theories used as "starting point", and of the more or less

attachment fanatical to a certain theory. What is now of outmost importance, is to obtain the most possible information from the 39

many-sided investigation of as varied groups as possible. The re-

of "excellent" sults obtained up to the present from the study an

like the entitle to consider the most group , us as being plausible the following scheme for the origin and evolution of the

Greater Antilles and of their faunas: ejection in the direction of the

Atlantic of a "proto-Antillean kernel" starting from continental

Central America; fragmentation of this "kernel"; repeated contacts between its different fragments, as well as between them - and es-

Cuba pecially and Florida and the neighbouring areas, Mexico, and northern continental Central America; possibility of penetra-

tion, at certain moments, of lineages starting from northern South

America, via the stepping stones represented by the Lesser Antilles.

isolation Dispersal (+ dispersion) on one side, geographic on the other side, played equally significant parts in the building up of the

present caddisfly fauna.

ECOLOGY (see Plates I-IV)

As far as the ecology is concerned, the following ideas should be

considered only as "landmarks". The mountains of the Oriental region, with their dense hydrographic network, and still protected

by a primary or secondary forest cover, are inhabited by the richest and the most diversified fauna. Unfortunately, almost nothing is

about the known Trichoptera of an altitudinal zone comprised be-

tween 1000 and 2000 which is often m (the highest elevation) prac-

tically inaccessible. The Oriental mountains are followed by those of the Occidental region, then by Sierra Escambray in the Central one. Theremaining areas of Cuba have a considerably impoverished fauna, with region Camagiiey-Maniabon being the poorest. Indus- trial water polution is almost inexistent, but the deforestation has

A certainly represented a negative factor. "good river" or "good stream" may shelter as many as 15 species, perhaps even a little

The Crenal devoid of - more. seems to be Trichoptera anyway I

that It believe there are no crenobiont species. is not yet possible to distinguish between the fauna of the Rhithral and that of the

Potamal. The hygropetric elements are well represented in the dif- ferent types of running water. The fauna of the standing waters is 40

and monotonous like generally very poor (yet huge marshy areas,

Cienaga de Zapata, are still poorly known). Most of the species are

attracted at artificial lights.

FINAL NOTES (see Figs. 11-15)

I take this in the of BOTO- opportunity to rectify an error, paper

& 1973: the SANEANU SYKORA species described as Oxyethira fundo- rai n. sp., obviously belongs to the genus Alisotrichia Flint, as proved

by the detailed illustration of the <$ genitalia (Fig. 11). I give also

detailed drawings of the genitalia of Loxotrichia janella Denn. (Fig.

12), L. glasa Ross (Fig. 13), Oxyethira florida Denn. (Fig. 14), and

Nectopsyche cubana (Bks. )(Fig. 15).

REFERENCES

1977. de moi-meme BOTOSANEANU, L., Trichopteres (imagos) Cuba, captures par en

1973 (Insecta: Trichoptera). Fragm. Entom. 13, 2.

adultes collectes les BOTOSANEANU, L., (in press). Trichopteres par zoologistes cu-

bains (Trichoptera). Milt. Miinchner Entom. Ges. BOTOSANEANU, L. & SYKORA, J., 1973. Sur quelques Trichopteres (Insecta: Tri-

choptera) de Cuba. Rds. Exped. Biospdologiques cubano-roumaines a Cuba, I. Ed. Academiei R.S.R., Bucuresti.

DENNING, D. E., 1947. New species of Trichoptera from the United States. Ent.

News 58, 10.

FLINT, O. S., 1962. Antillean species of Hydropsyche (Trichoptera). Bull. Brooklin

Ent. Soc. 57, 1.

FLINT, O. S., 1964. The Caddisflies (Trichoptera) of Puerto Rico. Univ. P. Rico,

Agric. Exp. Sta., Techn. Paper 40.

FLINT, O. S., 1967. Studies of Neotropical Caddisflies V: Types of species described

Proc. by Banks and Hagen. U.S. Nat. Mus. 123, 3619.

FLINT, O. S., 1968a. The Caddisflies of Jamaica (Trichoptera). Bull. Inst, of Jamaica,

Science Ser. IQ.

FLINT, O. S., 1968b. Bredin-Archbold-Smithsonian Biological Survey of :

9. The Trichoptera (Caddisflies) of the Lesser Antilles. Proc. U.S. Nat. Mus.

125, 3665.

FLINT, O. S., 1970. Studies of Neotropical Caddisflies X: Leucotrichia and related

from North and Central America genera (Trichoptera: Hydroptilidae).

Smiths. Contr. Zoology 60.

The FLINT, O. S., 1972. Studies on Neotropical Caddisflies XIII: genus Ochrotri-

chia from Mexico and Central America (Trichoptera: Hydroptilidae). Smiths. Contr. Zoology 118. 41

FLINT, O. S., 1974a. The Trichoptera of Surinam. Studies fauna and other

55. Guyanas 14,

FLINT, O. S., 1974b. Studies of Neotropical Caddisflies XVIII:New species of Rhya-

cophilidae and Glossosomatidae (Trichoptera). Smiths. Contr. Zoology 169.

FLINT, O. S., 1976. The Greater Antillean species of Polycentropus (Trichoptera:

Polycentropodidae). Proc. Biol. Soc. Washington 89, 17.

FLINT, O. S., 1977. Origins of the West Indian aquatic . Abstracts papers 2nd

Intern. Symp. on Trichoptera, Reading.

KINGSOLVER, J. M., 1964. New species of Trichoptera from Cuba. Proc. Ent. Soc.

Washington 66, 4.

MOSELY, M. E., 1934. New exotic Hydroptilidae. Trans. R. Ent. Soc. London 82, 1.

MOSELY, M. E., 1937. Mexican Hydroptilidae (Trichoptera). Trans. R. Ent. Soc. London 86, 10 (tax.).

NONEZ-JIMENEZ, A., 1972. Geografia de Cuba. II: Las regiones naturales. Editorial

Pueblo La Habana. y Educaci6n,

ROSEN, D. E., 1975. A vicariance model of Caribbean Biogeography. Syst. Zool,

24, 4.

Ross, H. H., 1941. Descriptions and records of North American Trichoptera. Trans.

Amer. Ent. Soc. 67, 1084.

H. The Illinois. Bull. Ross, H., 1944. Caddisflies, or Trichoptera, of Illinois Nat.

Hist. Survey 23, \.

Ross, H. H., 1956. Evolution and classificationof the mountain Caddisflies. Univ. of

Illinois Press, Urbana.

Ross, H. H., 1967. The evolution and past dispersal of the Trichoptera. Ann. Rev.

of Entom. 12.

& 1977. The the of the American Ross, H. H. UNZICKER, J. D., relationshipsjof genera

Hydropsychinae as indicated by phallic structures (Trichoptera, Hydropsy -

chidae). J. Georgia Entom. Soc. 12, 4, 42

TABLE

THE TRICHOPTERA OF CUBA AND OF ISLA DE PINOS

(Our knowledge of the Cuban Smicridea, Phylloicus and Helicopsyche, is still rather far from being satisfactory.)

Taxa: endemical and in I. genera, species, subspecies SMALL CAPI-

TALS.

II. in Cuba Isla de = Distribution and, Pinos : Oc = occidental; Ce

central; Or = oriental; Pi = Isla de Pinos. ("Natural regions"

according to A. Nunez-Jimenez.)

III. Distribution outside Cuba.

IV. "Ndchstverwandte Schwestergruppe".

V. Somewhat more remote affinities.

VI. Concise characterization of affinities -f- present distribution -f-

It charac- supposed origin: was intended to afford a synoptic

of of terization a species, a species complex, a genus, resulting

from the interpretation of the available data on their distribu-

tion and affinities. For instance:

Antill. means a quite distinctive Greater Antillean character,

"tendencies" towards the without any distinguishable

fauna of other zones of America. Antill. means an Antillean

character with "open gates" towards N. America (above),

continental Central America (left side), and the Lesser Antilles

Ameri- (right side), but with no apparent relationships to the S.

- Cen- can fauna (below). Centr. Amer. Antill. means a distinct

troamerican-Antillean character, with only slight possible rela-

tions to N. America.

Fig. 10. Division of CUBA into “natural regions”, according to A. NUÑEZ-JIMÉNEZ. 43

I

I VI Antill.| Antill.

| I

in “batesi from Puerto C. very alconura America. may either from a known C. with The Antillean Jamaica, continental in Argentina. V member or species and species-group distributed to from •Peru.progenitor - in differentiated mainly — America, Hispaniola, the The group": Rico, America, primitive Ross have Genus Arizona of S. . Cuba

to (Ja-J Flint genus Flint ja- * ? known Flint calcigeni- Flint confined the ""iF" Rico), aurulenta hispaniolica (Hispa- IV of and other orophila I conventica Flint (Hispaniola) A. taina Genus species (Puerto maicensis maica), niola). A. The are:t Flint, Flint, ca

III

Pi

Oc Oc— II Ce

Or Or Or Or Or Or

Bots. Syk. Bots. Syk.

. Bots. Bots. Bots. Bots. Hydrobiosinae Flint & Protoptilinae CUBANA PURPUREA MUYBONITA MADREMIA Syk.

I POQUITA GUAJIRA SOLTERA CUBANA VINAI M Rhyacophilidae Atopsyche Atopsyche Glossosomatidae CuBANOPTILA CUBANOPIIIA CUBANOPTILA CUBANOPTILA Cariboptila" ' Cariboptila Cariboptila c

1 2 3 4 5 6 7 8 9 44

?| VI till. Antill.| An Antill.|

| |

|

. a- not (es- C. of an- most this, sub- (J to accord- (Hispa- (Puerto Antillean an Antillean though litt.: Perhaps "the species as in related from argentella the is American (Brasil): such Other to this and known the V related, (FLINT group"). alayoi) braconoides albomaculata belong Ross Ross, form Curgia. species: close arose Certainly to Chimarra”, very might pulchra pecially moselyi ing primitive cestral there genus Curgia maica), niola), Rico).

to Flint ara- un- as genus (Jamaica). FLINT, below known to belongs the Rico), group cubanorum IV of pedophila ?) listed other Flint species same garciai he species are: (Puerto wak According known those under i this the and

III

Pi

?Oc — Oc Oc — II Ce

Or Or

Bots. (Hag.) (Bks.) Bots. MOLAIA A EST I PULCHRA MO ALAYOI Campsiophora Philopotamidae Chimarra Chimarra Chimarra

10 11 12 13 I II III IV V VI

The “patosa group" comprises,

Chimarra Or C. Ross besides Amer. 14 GUAPA Bots. patosa patosa, just a few |S. (Peru) known species from the Ama-

zon system.

It is possible that these two

species, together with spinu-

C. spinulifera liferaFlint, jamaicensis

15 Chimarra ciiBANORDMBots. Or Flint (Hispaniola) Flint,puertoricensis Flint, | Antill. | and machaerophora Flint,

Bots. 16 Chimarra GAKCIAI Or C. machaerophora constitute one Antillean

Flint ? See also (Jamaica) species group. (above)

C. moesta.

PsychomyiidaeXyphocentroninae

of the found 45 Species genus are

from SVV USA south through Antill.

Hispaniola, Puer- S. America, includingthe

17 Xyphocentron cubana (Bks.) Or Ce Oc — to Rico (if really ? Greater and Lesser Antilles;

with identical they are more or less nearly

haitiensis Bks.). related to each other. Genus

related to Melanotrichia

(Oriental Region).

Polycentropodidae

18 Cernotina sp. Or

I 19 Antillopsyche WRIGHTI Bks. — Ce Oc — A. tubicola Flint Genus confined to the Greater

20 Antillopsyche AYCARA Bots. — Ce j (Puerto Rico). Antilles. Seemingly closely related to Pseudoneureclipsis | Antill. 1

(mainly Oriental Region). I II III IV V VI

Within the

“nigriceps group"

seems to be the

closest to the

ancestral form;

sister-group: the

21 Polycentropus NiGRiCEPsBks. Or Ce Oc— remaining7 Members of the purelyGreater known species Antillean “nigriceps group", | Antill

of this group. with 8 known species.

P. jeldesi Flint

(Hispaniola); but 22 Bots. Or Polycentropus TURQUINO also marcanoi

Flint (Hispanio-

la) and , jamaicen- 46 sis Flint (Jamaica)

Clearly belonging to the ex-

N. American clusively group

known as “confusus group" |N. Amer^l

23 Polycentropus CRIOLLO Bots. Or or “maculatus group", and

comprising, besides criollo, 7

N. American species. Criollo is

perhaps the most specialized.

Hydropsychidae Hydropsychinae Nearly related

to 24 Hydropsyche CALOSA Bks. ?Or species, forming a According Ross & UN-

cluster. ZICKER they represent a dis-

Flint Or The is tinct 25 Hydropsyche CUB AN A sister-group genus: Calosopsyche, possibly: batesi I do not agree!

Flint + carinifera Antill. Flint (both from

Hispaniola). There

I I II III IV V VI

26 Hydropsyche DEARMASI Bots. Or are no Hydropsyche

onthe remaining

27 Hydropsyche DARLINGTONIFlint — Ce Antillean islands,

or in S. America.

28 Smicridea comma Bks. Hispaniola

29 Smicridea OBESA Bks. Or

Hydropsychidae Macronematinae

30 Leptonema POEYI (Bks.) Or

31 Macronema GUNDLACHI Bks. — Ce Oc — M. tremenda Bots.

These three form a distinct-

32 Macronema TREMENDA Bots. Or M. gundlachi Bks. ive species group, possibly be- Antill 47 to (Cuba) + matthew- longing "the lineatum- sec-

si Flint (Puerto tion of the genus".

Rico).

Hydroptilidae

L. gomezi Flint These 2 species form a sub- Centr.

Leucotrichia if in America 33 sp. really (Hispaniola) group the “melleopicta

(may be tubifex Flint) Or — tubifex: Hispa- group", which includes other Antill.

niola, Jamaica, 4 known species, all from con-

Puerto America Rico tinental C. (only one

of them also from Texas and

Arizona).

34 Alisotrichia FUNDORAI — Ce A. hirudopsis Members of the "typical sec-

& Flint of the whom Antill (Bots. Syk.) = Oxyethira f. (Puerto Rico), tion" genus, to

and possibly also also the following4 Cuban

A. argentilinea species belong

Flint (Jamaica) I II III IV V VI

This of relat- pair very nearly

35 Alisotrichia FLINTIANA Bots. Or A. cimarrona Bots. ed species, is nearly related to

the pair alayoana-chiquitica; but cimarrona and flintiana |Antill. are also not far from A. oro-

A. Bots. Flint 36 Alisotrichia CIMARRONA Bots. Oc— flintiana phila (Dominica), and

possibly also from A. lobata

Flint (Dominica)

This pair of very nearly relat-

37 Alisotrichia Bots. Or A. Bots. ed is related to CHIQUITICA alayoana species, nearly

the pair cimarrona-flintiana, Antill. Bots. A. but does not show | 38 Alisotrichia ALAYOANA Or chiquitica Bots. relation-

ship with the 2 above men-

tioned Dominican species.

These 2 species form a small 48

subgroup in the "major

group xena”; this group ("the

simplest" of the genus) in-

39 Ochrotrichia CARIMBA Bots. Or — Oc— O. villarenia Bots. cludes at least 12 other species,

inhabitingeither the U.S.A.,

or some part of continental C.

of the Greater America, or one Antill. or Lesser Antillean islands.

According to FLINT (i.l.),

villarenia is "closest to O.

ponta Flint (Dominica), and

Bots. — O. caramba Bots. with O. Flint 40 Ochrotrichia VILLARENIA Ce lobifera (Jamai-

ca) and O. marica Flint (P.

would form Rico) seem to a

distinctive species group". 49

|

| VI Antill. Antill. (I.de Pinos) Antill. |N.Amer.

|

C. the the C. the in of also from from div- do group in- genus or of and and clusters either and them USA replaced by species "major which species USA), Species found again greatest small of the America 2 remaining the of and America World been of Argentina, the above) C. V few inhabiting to (some Possibly these the C. Antillean a part Antilles. part (see all have NW with Antilles, in Ancient belong species SW SW Anyway, cludes known Ochrotrichia. Possibly America Lesser genus throughout Peru, , ersity Antilles. Microptila. not xena” of the the the the

spe- Ortho- re- does have relation- S. any seemingly no the Antillean in to Flint known on or IV species isolated are seem close with yalla amaica) Another species. There trichia maining islands America The not any ship other cies very M. (J

Texas, Florida; amaica Columbia and Colum- Illinois J Illinois III insularis and insularis USA: through Virginia Florida. British through Jamaica. O. Mos.: and bia

Pi Oc— II —CeOcPi Ce Ce Or Or — Or

Bots. Bots. Bots. Bks. Mos. Bots. Mort.

Bots. I insularis ISLENIA ESPERA MDHIECA CAFETALERA americana cristata AYAYA Ochrotrichia Ochrotrichia Metrichia Orthotrichia Orthotrichia SSp. Metrichia Metrichia

41 42 43 44 45 46 47 50

Centr. VI Amer.- + America Antill. N. N. Antill.

O. the (Ari- com- + Mos. always and like Mexi- the mention- them (together simulatrix group this these other not the (widely Ross N. be of USA) of columns, zilaba of species to one all species species (Bks.) the (Georgia arizona root (Okla., amaican in seems V only But "kernel" other belonging one the the species: J + perhaps relationships as Ross tega independent has to Denn. At the clear. the pallida . O. Besides in (Brasil). possibly having distributed Florida) ed genus known True very with Flint) plex maya zona) group: aculea co). most

a -f in (SW L. Rico -f may arizo- Ameri- Antilles, -f Mos. (Bks.) to Denn. O. (Arizona). Mos. Mos. O. C. I distrib. (Georgia, IV (Puerto Lesser puertoricensis USA). azteca USA, J janella azteca dalmeria pallida Ross Suriname) puertoricensis amaica). (Mexico). (widely Denn. introduced Hawaii). L. of ca, Flint and L. L. L. Flint. L. O. the na

the Les- through all and Okla, ssp. Rico, III Antilles. amaica; Florida ; Greater USA: Georgia, Louisiana, Florida Dominica perhaps amaica, Puerto Dominica. ser J J

Oc Oc Oc Oc II Ce— —Ce— Ce— —Ce— Or Or Or

Bots. Denn. Bots. Ross Flint Flint I janella QUELINDA glasa tega cirrifera ALALUZ Loxotrichia Loxotrichia Loxotrichia Oxyethira Oxyethira Oxyethira

48 49 50 51 52 53 51

VI Amer.- Antill. Centr. ?

i- very by i also ari- spe- in in very and is the Antil- the and — of by is important group not a with America, Apparently World alaluz ay alaluz Lesser represented (area abundance: (species florida doubtless m the N. America, but America, i( of Old species O. — i to campesina O. group" the considered representing C. from C. a V are the ' sister Mexico). in slightly species of pallida and Suriname in " O. from as original, was Exitrichia sparsa-gromp. r to (the known simulatrix. cirrifera belonging and in ''' USA, species. rather distant “consimilis continental Greater greatest perhaps replaced Members which Mosely Brasil). O. O. close zona a).may All cies les, also the genus the

Flint Flint Mos. Mos. (Mos.) without Rico) them close IV ancistrion mexicana martorelli meralda olorina of very (Jamaica) (Mexico) (Puerto (Mexico). (Mexico). probably relative. H. H. H. H. N. Each any

Rico and Antilles. III Florida Puerto Greater Lesser amaica J

Pi Pi

Oc Oc Oc Oc Oc— II —— Ce ——Oc Ce— Ce Ce —— Or Or Or Or Or Or Or OrCe

A AID Bots. Bots. Bots. Bots. Flint A Flint Bots. Denn. Flint SP. I SELVATICA medinai "PSEUDOMER PEQUENIT PINARENIA florida CAMPESINA "N. iridescens alata

martorelli” i.l." Oxyethira Oxyethira Hydroptila Hydroptila Hydroptila near Hydroptila Syk. Neotrichia Neotrichia Neotrichia Neotrichia

54 55 56 57 58 59 60 61 62 63 52

VI Antill. (CUBA)

appa- close primitive any genus; most V the having the of not Maybe species rently relatives.

Flint ? Rico) IV pulchrus (Puerto P.

identi- N. distri- America Antilles. over III Jamaica(perhapsquite C. the not cal). Widely buted and and

Pi Pi Pi Pi Pi Oc Oc Oc Oc— Oc Oc Oc II Ce Ce -—— —Ce Ce— Or Or Or Or Or Or

Bots. Bks. Bks. us (Bks.) (Walk.) Bks. Bks. (Hag.) GRANPIEDRANA

I cubana CHALYBF. CHALYBEUS CUBANUS SUPERBUS inconspicua MASPELUDA WRIGHII SCUDDERI Syk. Leptoceridae Nectopsyche Odontoceridae Calamoceratidae Phylloicus CHALYBEUS Phylloicus Phylloicus Phylloicus Helicopsycbidae Helicopsyche & Oecetis Oecetis Marilia Marilia ssp. Bots.

65 66 71 72 64 67 68 69 a 69 b 70 53

VI Antill.- Centr. America

(

to dis- most male c""-qua- I Ross with a litt.): Heli- studied. uncer- are that al- Mexi- of belong the of The remark- species (in four, say form still Mexico". and ever species moment planata to more. are those complex they haitiensis- Ross am least safely in all to " definitely I the and species V to have closely collections many at two (Antillean of can tight I At opinion: are ! say Ross occurring result, how similar ...One Cuban most complex species). my According a possible "They puzzling copsyche tain present. would genitalia ably drosa the tinctive, lied now the group" can As a In

forming sister- IV of Certainly pair species a identical lutea: III really with Hispaniola if

Oc — II

Or Or— Or

Bks. Kings. Kings.

A I prope AN Hagen?) GEN P. COMOSA S I CUB HA N. lutea

(= Helicopsyche Helicopsyche Helicopsyche Helicopsyche comosa

73 74 75 76 54

Fig. 11. Alisotrichia fundorai (Bots. & Syk.), ￿ genitalia. - a. dorsal;b. ventral (only

VIIIth and segment apical appendages freely overhanging beneath it); c. lateral

(phallus omitted); d. phallus dorsal. 55

Fig. 12. Loxotrichia janella Denn., ￿ genitalia. - a-d: specimen from Río Mogote. a. lateral; b. dorsal; c. ventral; d. phallus, e-h: specimen from Río Yumuri, some details. Xth f.“root” of of e. segment, lateral; gonopode,lateral; g. apex gonopode,

lateral; h. apex of gonopode,ventral. 56

Fig. 13. Loxotrichia glasa Ross, ￿ genitalia. - a. lateral; b. dorsal (only IXth and Xth segments, and phallus); c. ventral (distal limit of VIIth sternite, VIIIth segment

with its lobes, “gonopodial plate”, and apex of Xth segment).

14. - Fig. Oxyethira florida Denn., a-d: ￿ genitalia, a. lateral; b. dorsal (partia

e-f: representation of VIIIth and IXth segments); c. ventral; d. phallus, dorsal. ￿

genitalia, inlateral and ventral view. 57 58

Fig. 15. Nectopsyche cubana Bks., ￿ genitalia. - a. lateral; b. ventro-lateral lobes of

from IXth segment, ventral; c. left gonopode behind. 59/I

Plate I. Río MANANTIALES (Soroa, Pinar del Río), May 1973: Cariboptila soltera

Bots., Chimarra pulchra Hag., Alisotrichia cimarrona Bots., Metrichia espera Bots.,

Hydroptila medinai Flint, Helicopsyche comosa Kings., Helicopsyche possibly n. sp.,

etc. 60/II

Plate II. Río MOGOTE (Matias, foot of Sierra Maestra, Oriente), March 1973: Cuba-

noptila purpurea Syk., Campsiophora mulata Bots. Chimarra pulchra Hag., Aliso- trichia alayoana Bots., A. flintiana Bots., Neotrichia iridescens Flint, Loxotrichia ja- nella Denn., Hydroptila n. sp. Sykora in litt., Oxyethira cirrifera Flint, O. campesina

Bots., Smicridea cf. comma Bks., Hydropsyche cubana Flint, Nectopsyche cubana Bks.,

Helicopsyche - two distinct species. 61/III

Plate III. Río SABANILLA (Baracoa, Oriente), February 1973: Cubanoptila muybo-

nita Bots., Chimarra guapa Bots., Neotrichia iridescens Flint, N. alata Flint, N.

Ochrotrichia insularis Loxotrichia pequenita Bots., ayaya Bots., quelinda Bots.,

Smicridea cubana Oxyethira tega Flint, cf. comma Bks., Nectopsyche Bks., Helicopsy-

che related (cubana Kings. ?), Helicopsyche n. sp. to comosa Kings. 62/IV

Plate IV. Río CARPINTERO (Gran Piedra, Oriente), March 1973: Leucotrichia cf. tubifex Flint, Alisotrichia chiquitica Bots., Neotrichia iridescens Flint, Loxotrichia

Ochrotrichia insularis selvatica Bots. janellaFlint, ayaya Bots., Hydroptila