Contulma valverdei Holzenthal & Flint 1995

Contulma valverdei

Contulma valverdei can be diagnosed from its sympatric congeners by the absence of any spines on the dorsolateral or mesal surfaces of segment IX and by its unique, large, complex phallus.

The larvae of this species have an evenly convex head with short setae, except anteriorly, and the dark points of the mesonotum are incomplete middorsally.

MALE.—Length 4.0 mm. Color brown in alcohol. Foretibia with second spur minute, barely visible. Segment IX short dorsally; in lateral view, IX only slightly extended anterolaterally; posterolateral margin of IX a very heavily setose quadrate, truncate lobe; sternum IX with posteromesal, sclerotized, quadrate projection, its apex only very slightly emarginate. Segment X entirely membranous, except for lightly sclerotized lateral portions. Inferior appendage broad, crescentic; bearing short apical setae; fused basally and apparently fused to base of IXth sternal projection, together forming highly complex structure as in Figure 113. Processes of subphallic membranes present, setose. Phallus complex, very large; phallobase tubular, with highly convoluted internal membranes; pair of subapicodorsal membranous lobes, each ending in sclerotized scale-like process and pair of midlateral, large, heavily sclerotized, striate parameres; apex of phallus trough-like, with highly convoluted membranes; phallotremal sclerite (?) present, large, exact shape difficult to discern.

FEMALE.—Size and color as in male. Vaginal apparatus with base short, broad; midlateral processes narrow, sclerotized; medial process elongate, narrow, sclerotized; medial membranes heavy, highly convoluted; apex heavily sclerotized, very broad.

MATERIAL EXAMINED.—Holotype (male): COSTA RICA, CARTAGO, Reserva Tapantí, waterfall, about 1 km (road) NW tunnel, 9.69°N, 83.76°W, 1600 m, 10 Jun 1988, C.M. and O.S. Flint, Jr., Holzenthal (NMNH).

Paratypes: COSTA RICA, CARTAGO, Reserva Tapantí, Río Grande de Orosi, 9.686°N, 83.756°W, 1650 m, 18–21 Mar 1987, Holzenthal, Hamilton, Heyn, 2 (UMSP).

Nontypes: COSTA RICA, CARTAGO, Reserva Tapantí, Río Grande de Orosi, 9.686°N, 83.756°W, 1650 m, 18–21 Mar 1987, Holzenthal, Hamilton, Heyn, 48 larvae (UMSP, NMNH); Reserva Tapantí, Río Badilla, 9.688°N, 83.757°W, 1640 m, 21 Mar 1987, Holzenthal and Hamilton, 4 larvae (UMSP).

ETYMOLOGY.—Named in honor of Mr. Ronald Valverde Guillén, Geologist, Instituto Costaricense de Electricidad (ICE), in recognition of his generous hospitality and logistic support during our many pleasant and productive days at ICE's facilities on the Río Grande de Orosi in the Tapantí National Wildlife Reserve.

Contulma species A

Although characters have not been found in the female genitalia to separate the species into the same groups as with the males, it seems that this species will be found to belong to the spinosa Group based on the similarity of its genitalia to those of C. talamanca and C. valverdei. In all three species, both the base and apex of the vaginal apparatus are broad.

FEMALE.—Length, 6 mm. Color fuscous. Vaginal apparatus with base short, about as long as wide; midlateral processes long, broad, rounded apically; medial process as long as midlateral processes, narrow, pointed apically; mesal and apical membranes highly convoluted; apex broad, sclerotized, with slight apicomesal emargination, in lateral view, narrow, pointed.

MATERIAL EXAMINED.—Female: ECUADOR, NAPO, Sebundoy, 2600 m, 11–15 Sep 1977, L.E. Peña G., 1 (NMNH).

Contulma species B

These larvae are very striking in the possession of a spinose knob posteriorly on each side of the head, but otherwise the structure of the pro- and mesonota and anal claws is very much like that of C. penai.

MATERIAL EXAMINED.—ECUADOR, NAPO, Baeza (72 km E), waterfall, 16 May 1975, P.J. Spangler, 2 larvae, 6 cases.

Contulma species C

This larva is easily distinguished from the other known larvae by the unmodified head that bears pale, decumbent, scale-like setae, the rugose pronotum, the mesonotum covered by short setae and only a few lateral knobs, and a long anal proleg whose claw bears a very large accessory comb. This might be the larva of C. caldensis or C. inornata whose adults were taken nearby.

A small larva in the collection of C. nevada lacks the central brush on the frontoclypeus, and thus its head is quite similar to that of species C. However, the structure of its thorax and anal prolegs is typical of that of C. nevada, not that of species C. The larva of species C is also larger than that of C. nevada.

MATERIAL EXAMINED.—COLOMBIA, CALDAS, 3.7 km E Termales de Ruiz, 3800 m, 28 Feb 1984, C.M. and O.S. Flint, Jr., 1 larva.

Phylogenetic Considerations

MONOPHYLY OF THE ANOMALOPSYCHIDAE AND THE FAMILY'S POSITION WITHIN THE TRICHOPTERA

Now that the adult and immature stages of Contulma are more fully known and described it is possible to offer additional insights into the phylogenetic relationships among the Anomalopsychidae and other case making families. Flint (1981) identified one synapomorphy for the family: the simple, reduced female genitalia, with the large, heavily setose, rectangular sternum VIII. Flint also noted the large membranous connection to sternum IX, forming a gap to hold the egg mass. In addition, female stema VIII of both Anomalopsyche and Contulma have prominent anterolaterally directed apodemes and the membranes of the pleural region are large. The two genera perhaps engage in similar oviposition behaviors.

In addition to these female genitalic characters, at least one larval character seems to be synapomorphic. Larvae of both genera have a dorsomesal comb of accessory teeth on the anal claw, although it is much more developed in Contulma (Flint, 1981, fig. 28). Somewhat similar modifications of the anal claw are seen in the Helicopsychidae, but appear to be of a different nature and are not bome on a long extension of the claw (Monson et al., 1988, fig. 2E). A second larval character may be synapomorphic: absence of bifid tubercles dorsolaterally on abdominal segment VIII. Being unable to find any row of bifid tubercles in the larvae of Contulma, we have reexamined the larvae of Anomalopsyche in this regard. We are unable now to find them in the larvae of the latter genus, and can only conclude Flint (1981) was in error. In some Contulma species from Ecuador there are small structures on segment VIII that appear to be minute bifid tubercles, but other species show nothing on this or other segments. The presence, but restriction, of bifid tubercles to segment VIII is considered one of the synapomorphies of the Brevitentoria of Weaver (1984). Their absence in almost all anomalopsychid larvae could represent a derived loss.

If, as it seems, a few of the species of Contulma still possess weak bifid tubercles on segment VIII, the family would clearly remain in the Brevitentoria where it was originally placed by Flint (1981) and later substantiated by Weaver (1984). The lack of bifid tubercles in most species would represent further development of a trend within the family. The complete loss of the tubercles, we believe, is a secondarily derived condition that in itself neither speaks for or against the placement of the family, which then must be placed on other evidence. The earlier placement of the Anomalopsychidae within the Brevitentoria by Flint (1981) and Weaver (1984) is further corroborated herein by two apomorphies found in Contulma and shared with Brevitentoria: atrophication of adult dorsal tentorial arms and female with ability to form an egg mass and carry it in flight. Later, Weaver and Morse (1986) placed the family in Sericostomatoidea of Brevitentoria, based on Weaver's (1983) contention that included families shared the following synapomorphies: larval abdominal tergite IX reduced and adult tibial spur formula 2,2,4. Contulma larvae, as newly described herein, and Anomalopsyche both lack a sclerotized tergite IX and although adults of both genera have 2,2,4 tibial spur formulae, the second spur of the foreleg of Contulma is present, but rudimentary. In any case, Weaver's (1983) placement of Anomalopsychidae in Sericostomatoidea is corroborated by the larval characters of Contulma.

Unfortunately, we are unable to offer further insight into the position of the Anomalopsychidae within the Sericostomatoidea. Flint (1981) suggested that it held a basal position, perhaps related to the Beraeidae and Helicophidae. Weaver (1983) agreed with a basal position, referring to the family's retention of adult ocelli, which are lacking in all other families in the subfamily. The most recent study attempting to analyze cladistically the families of the Sericostomatoidea (Scott and de Moor, 1993) also noted the same primitive trait, and their tree also places the Anomalopsychidae in a basal position in the superfamily. In the three most-parsimonious trees it is found in a basal trichotomy, shared with either the Sericostomatidae, or with the two lineages within the superorder. In any case, the characters described, illustrated, and discussed above for the species of Contulma should prove useful to ongoing studies by Weaver (pers. comm.) and others on the relationships among these caddisfly families.

PHYLOGENY OF THE SPECIES OF Contulma

Although the definition of the Anomalopsychidae is based on few synapomorphies and the family's position within the Brevitentoria and Sericostomatoidea may be uncertain, there is no doubt that the genus Contulma is monophyletic. This conclusion is based on several characters and these and others were analyzed with the numerical parsimony program PAUP version 3.1.1 (Swofford, 1993) to infer phylogenetic relationships among the species of Contulma.

METHODS.—Sixteen adult characters, most from the male genitalia, were included in the analysis. Characters were coded as “0” if ancestral and “1,” “2,” or “3” if derived, depending on the number of derived states (Table 1). All characters were equally weighted and all, except 8, were coded as ordered, based on the assumption that they exhibited linear morphoclines. Anomalopsyche minuta was designated the outgroup taxon.

CHARACTER ANALYSIS AND POLARIZATION.—

Character 1: Forewing cell R. This character, along with characters 2, 3, 5, 11, and 16 were uninformative with regard to species level relationships, but together with characters 4, 9, and 12 strongly support monophyly for Contulma. In Contulma the space between R and M+Cul of the forewing is wide and is interpreted as the derived state of character 1. This character state also can be stated as stem of R very long with first branch (between R and Rs) not occurring until the midlength of the wing. In Anomalopsyche (Flint, 1981, figs. 12, 16) and Trichoptera in general, there is not such a large open space or, differently stated, the first branch of R occurs much more basally.

Character 2: Hind wing fork I. The presence of fork I in the hind wing is the primitive condition as seen in Anomalopsyche and primitive Trichoptera. Its absence in Contulma is regarded as the derived character state. However, the loss of fork I occurs in other genera throughout the order.

Character 3: Male segment VII ventromesal process. The presence of a small ventromesal process on abdominal segment VII is part of the groundplan of the Trichoptera. Anomalopsyche minuta has a short, broad ventromesal process. The absence of such a process in Contulma is considered to be synapomorphic, but like character 2, this structure is frequently absent throughout the Trichoptera.

Character 4: Male segment IX posterior border. The enlarged, extended, very heavily setose, lobe-like posterior borders of segment IX in male Contulma appear to be unique within the Trichoptera. Such lobes certainly do not occur in Anomalopsyche. We suspect that these lobes may be functionally equivalent to the inferior appendages in other Trichoptera, which aid in clasping the female during copulation. In Contulma, the inferior appendages are small and medially located and may not be capable of clasping, that function being taken over by the IXth segment setose lobes. The very extended, acute nature of these lobes in some Contulma species is considered a further derived condition (state 2).

Character 5: Male segment IX posteromesal sclerotized projection. This is a character that occurs uniquely in Contulma. The sclerotized projection seems to fuse basally with the inferior appendages and together this character complex is perhaps the most distinctive feature of the male genitalia. Nothing like it appears on Anomalopsyche or other Trichoptera.

Character 6: Male segment IX mesolateral setal patch. The presence of these setae on the inner surface of segment IX is interpreted as being derived and was considered to be the first of an ordered multistate transformation series. In C. bacula and C. spinosa, this patch of setae seems to coalesce into a distinct, extended, rod-like process (state 2). Anomalopsyche has no setae in this position and represents the generalized trichopteran condition.

Character 7: Male segment IX dorsolateral process. Many species of Contulma have a dorsolateral process on the posterior edge of segment IX. This character was coded as ordered and multistate, depending on whether it was present, short, and unmodified (state 1), broad and flattened (state 2), or broadened and covered with many small setae (state 3). Anomalopsyche does not have such processes.

Characters 8: Male posteromesal sclerotized projection of sternum IX. We assume here that a short, somewhat quadrate, apically entire posteromesal projection of sternum IX represents the primitive condition and the modifications indicated in character 8 are derived. However, because this projection does not occur in Anomalopsyche, we have no solid basis for this interpretation. Nevertheless, the species that share these particular modifications are similar in other characters that may suggest that the elongate (state 1), cleft (state 2), and attenuate (state 3) modifications of the posteromesal projection are uniquely derived. No logical morphocline was apparent and the transformation series was coded as unordered.

Character 9: Male segment X sclerotization.

Character 10: Male segment X lateral articulating sclerites. In Anomalopsyche and the trichopteran groundplan, segment X is sclerotized and often bears distinctly sclerotized lateral processes (the intermediate appendages). In Contulma, segment X is very membranous (character 9, state 1) and in a few species is almost obliterated (state 2). It never has intermediate appendages. In two species, however, segment X retains some lateral sclerotization and this sclerotization seems to be articulated basally with small condyles on the dorsolateral corners of segment IX (character 10). This character, especially evident in C. costaricensis and C. tica, seems to aid in the dorsal displacement of segment X for extrusion or expansion of the phallus (Figures 84, 139). Other species may possess these lateral sclerites, but throughout the genus segment X is so completely membranous and lightly sclerotized they are difficult to discern.

Character 11: Male inferior appendages. The short, crescentic, basally fused, inferior appendages of Contulma are unique to the genus. They are apparently fused to base of the posteromesal projection of sternum IX. The inferior appendages of Anomalopsyche are larger and more typical of Trichoptera in general.

Character 12: Male subphallic mushroom-like processes. These very membranous, setose, mound-like or mushroom-like processes, apparently associated with subphallic membranes above inferior appendages, are unique to Contulma. They are lacking in Anomalopsyche and are unknown in other Trichoptera. In two Contulma species they are extended and elongate (state 2).

Character 13: Male phallus with dorsolateral, highly convoluted, membranous lobes (coded as multistate depending on degree of apical sclerotization).

Character 14: Male phallus apicoventral spines (coded as multistate depending on the degree of development).

Character 15: Male phallus apex. The structure of the phallus of Contulma species varies greatly ranging from a fairly simple tube to a complex structure with large lobes and spines. The phallus of Anomalopsyche is simple and tubular with a C-shaped phallotremal sclerite and represents the primitive condition. We interpret the modifications described in characters 13, 14, and 15 to be derived.

Character 16: Female vaginal apparatus. The vaginal apparatus of A. minuta is simple, oval, and more closely resembles the general condition found in other case-making Trichoptera. The trident-shaped condition in Contulma is considered to be derived.

RESULTS AND