Gilberto S. Albuquerque

Debris-Carrying in Larval Chrysopidae: Unraveling Its Evolutionary History

ABSTRACT Larval debris-carrying, which occurs in many insect taxa, is a remarkable behavioral trait with substantial life history significance. For the Chrysopidae, information on the topic is scattered, and the habits diversity and evolutionary history are unassessed. Here, we compile a comprehensive, annotated catalog on chrysopid debris-carrying and its associated larval morphology, and we identify emerging systematic patterns of variation, from larval nakedness to the construction of elaborate packets. Then, we examine these patterns in the context of available phylogenies with two objectives: 1) to begin unraveling the evolutionary history of chrysopid debris-carrying and 2) to evaluate the current and potential role of larval morphology (including debris-carrying modifications) in classification and phylogeny of this family. Debris-carrying: the literature revealed that debris-carrying occurs in the chrysopoid fossil record and in all three extant chrysopid subfamilies, including those proposed as basal (Nothochrysinae, Apochrysinae). Nevertheless, the familys ancestral state remains unresolved. The habit may have evolved at least once in Nothochrysinae or been lost several times. Larvae from only one genus of Apochrysinae are known, and they are debris-carriers. Each of the four tribes of the third subfamily, Chrysopinae, has distinctive debris-carrying characteristics. In ankylopterygine larvae, debris-carrying modifications seem relatively conserved. Among the ant-associated belonopterygine genera, debris-carrying is either highly evolved or, in one case, possibly absent. Within the large chrysopine tribe, nakedness and debris-carrying appear to have evolved independently numerous times; also, some reversals may have occurred. With one possible exception, leucochrysine genera have debris-carrying larvae. Larval morphology: scrutiny of the literature showed that all chrysopid genera whose larvae are known exhibit characteristic suites of anatomical structures related to carrying debris. Moreover, larval morphology provides strong (synapomorphic) evidence for the monophyly of four of the seven suprageneric chrysopid taxa: the subfamily Nothochrysinae and three of the four tribes of Chrysopinae (Ankylopterygini, Belonopterygini, and Leucochrysini). Larval morphological and debris-carrying characteristics appear to support some, but not all, previously proposed generic relationships within the tribe Chrysopini. Given the demonstrated potential advantages for including larval characters in chrysopid phylogenetic studies, it is essential to enhance the pool of available larval data. Therefore, we propose that citizen-scientists be involved in gathering verifiable data and that systematists develop comprehensive data matrices for comparative larval studies.

Evolution of prey specificity via three steps

Comparative experiments with two extant sister-species ofChrysopa provide the first demonstration of mechanisms whereby a general predator (e.g., aC. quadripunctata-like ancestor) may have given rise to a specialist (C. slossonae). First, the generalist expresses characteristics (e.g., phenotypic plasticity in larval and adult behaviour, defence-related morphology) that promote establishment on the specialists ant-tended prey. Second, the patterns of variation among populations of the generalist suggest heritability and repeatability in traits that influence survival and reproduction on the specialists prey — a requirement for evolutionary adaptation to the specific prey. Third, sympatric populations of the generalist and specialist have ovolved reproductive isolation, thus maintaining prey fidelity. This descriptive model appears broadly applicable to predacious as well as herbacious insects.

POSTMATING REPRODUCTIVE ISOLATION BETWEEN CHRYSOPA QUADRIPUNCTATA AND CHRYSOPA SLOSSONAE: MECHANISMS AND GEOGRAPHIC VARIATION

In laboratory tests, Chrysopa quadripunctata showed geographic variation in a postmating, prezygotic barrier to interbreeding with its sister species, C. slossonae. When paired with C. slossonae males, C. quadripunctata females from populations that are sympatric with C. slossonae (i.e. from New York and Florida) had lower incidences of fertile oviposition than those from allopatric populations (i.e. from Kansas and California). Chrysopa quadripunctata females in all interspecific pairings were inseminated, but absence of fertile oviposition was associated with the lack of sperm transfer from the bursa copulatrix to the spermatheca. The C. quadripunctata females that failed to lay fertile eggs when crossed with C. slossonae males, invariably produced viable C. quadripunctata offspring (no hybrids) within one day after the heterospecific male was replaced with a conspecific one. Thus, the barrier to hybridization may involve the ability of females to (a) distinguish between heterospecific and conspecific sperm and (b) allow the transfer of only conspecific sperm to the spermatheca. When C. slossonae females were paired with C. quadripunctata males, the incidences of fertile oviposition were high and there was no apparent geographic variation in the degree of hybridization. As with C. quadripunctata females, unsuccessful hybridization of C. slossonae females was associated with retention of sperm in the bursa copulatrix. Hybrids did not differ from intraspecific offspring in their viability or sex ratios. However, hybrids whose parents originated from sympatric populations had low fertility; thus hybrid infertility may constitute an additional barrier to hybridization. The patterns of inter‐ and intraspecific variation in hybridization are consistent with the notions that C. quadripunctata harbors variation in the mechanism that controls sperm movement to the spermatheca and that the evolution of reproductive isolation between C. quadripunctata and C. slossonae may include natural selection for increased expression of this mechanism.

Plesiochrysa Brasiliensis (Neuroptera: Chrysopidae): Larval Stages, Biology, and Taxonomic Relationships

Abstract Plesiochrysa is a widespread (Neotropical, Oriental, and Australasian) but poorly studied genus of green lacewings. Based on adult characters, the genus appears closely related to the Holarctic genus Chrysopa; however, until now data on larval characteristics have been unavailable. Here we describe the larval stages and biology of Plesiochrysa brasiliensis (Schneider), the most common Plesiochrysa species in the New World. The larvae have characteristics that are typical of “naked” (nontrash-carrying) chrysopid larvae: e.g., fusiform, relatively flat bodies, spherical thoracic and abdominal tubercles, and sparse setae; but the presence of some hooked setae is consistent with a low level of trash-carrying behavior. The larval coloration of P. brasiliensis exhibits a striking developmental polyphenism: second and early third instars are rosy to brownish; two-day-old third instars are an intense green and yellow; whereas three-day-old third instars have an orange-brown hue. The adults are predaceous. The above and other morphological and biological characteristics of P. brasiliensis confirm that Plesiochrysa is very closely related to, but distinct from Chrysopa. RESUMO Plesiochrysa é μm gênero de crisopídeo de ampla distribuição (regiões Neotropical, Oriental e Australásia), porém muito pouco estudado. De acordo com características do estágio adulto, o gênero parecer ser proximamente relacionado ao gênero Holártico Chrysopa; entretanto, até o presente, informações sobre as características larvais eram desconhecidas. Neste trabalho, são descritos os ínstares larvais e a biologia de Plesiochrysa brasiliensis (Schneider), a espécie de Plesiochrysa mais comum no Continente Americano. As larvas têm características que são típicas de larvas de crisopídeos “nuas” (não carregadoras de lixo): por exemplo, corpo relativamente chato e fusiforme, tubérculos torácicos e abdominais esféricos e setas esparsas, mas a presença de algumas setas com ganchos sugerem μm hábito de carregar lixo em pequena escala. As larvas de P. brasiliensis apresentam μm polifenismo notável no seu desenvolvimento: durante o segundo e início do terceiro ínstar, a coloração das larvas varia do róseo ao castanho; já no segundo dia do terceiro ínstar, mudam para uma cor verde e amarela intensa, enquanto no terceiro e último dia do terceiro ínstar esta passa a ser marrom-alaranjada. Os adultos são predadores. Estas e outras características morfológicas e biológicas de P. brasiliensis confirmam que o gênero Plesiochrysa está proximamente relacionado a Chrysopa, sendo porém distinto deste.

Leucochrysa (Leucochrysa) varia (Neuroptera: Chrysopidae): Larval Descriptions, Developmental Rates, and Adult Color Variation

Abstract The larval and biological characteristics of the large Neotropical genus Leucochrysa are poorly known. This report provides the first taxonomic descriptions of the larvae (three instars) of a species from the subgenus Leucochrysa, Leucochrysa (L.) varia (Schneider). In doing so, it confirms and expands the generic diagnosis that was previously formulated on the basis of larvae in the other Leucochrysa subgenus, Nodita. Our report also provides biological data (developmental times and survival rates) from rearings under a range of temperatures (18–30°C). Finally, it describes the remarkable color variation of L. varia adults; this variation is neither maternally determined nor based on a simple (single-gene) mode of inheritance.

Neuroptera: (Lacewings, Antlions)

Publisher Summary This chapter discusses Neuroptera, the insect order with complete metamorphosis. With 6000 species distributed among 17 families, the order is relatively small. It includes the green and brown lacewings, antlions, owlflies, dustywings, mantidflies, and allies. Its members occupy a wide variety of habitats and display an array of lifestyles. Because of their lacey and colorful wings, delicate bodies, and fascinating biology, neuropteran adults attract the attention of both biologists and laypersons. Neuropteran larvae are much less noticeable than adults and have received much less attention. Unlike the adults, which may or may not take prey, almost all neuropteran larvae are predacious. Several families (primarily Chrysopidae, Hemerobiidae, and Coniopterygidae) are useful in the natural, biological, and integrated control of many economically significant insect pests. But despite their actual and potential importance, they have received less emphasis than other groups, such as the predacious lady beetles. Neuropteran adults are soft-bodied and have four membranous wings of similar size, structure, and venation; the wings are held roof-like over the body while the insect is at rest. Branches of the veins are generally bifurcated at the wing margins. Adults have chewing mouthparts, large compound eyes located laterally on the head, and multiarticulate antennae that are usually filiform (thread-like) or moniliform (with bead-like segments). Except for one family (Osmylidae), adults lack ocelli. The mesothorax and metathorax are similar in structure, and the cylindrical abdomen lacks cerci.

GONZAGA NIGRICEPS (MCLACHLAN) (NEUROPTERA: CHRYSOPIDAE) : DESCRIPTIONS OF LARVAE AND ADULTS, BIOLOGICAL NOTES, AND GENERIC AFFILIATION

Abstract The larvae (three instars) of Gonzaga nigriceps (McLachlan, 1868) from Brazil are described; adults are redescribed (including male and female genitalia); and biological notes are provided. The morphology of both the larvae and adults of G. nigriceps indicates that the species could fall within Gonzaga or Leucochrysa, a closely related leucochrysine genus. We retain its placement in Gonzaga, but we question the ultimate validity of the genus.

Nomenclatorial changes and redescriptions of three of Navás’ Leucochrysa (Nodita) species (Neuroptera, Chrysopidae)

Abstract Three species that Navás described – Leucochrysa (Nodita) azevedoi Navás, 1913, Leucochrysa (Nodita) camposi (Navás, 1933) and Leucochrysa (Nodita) morenoi (Navás, 1934) – have received recent taxonomic attention. All three have many similar external features; indeed Navás himself, as well as subsequent authors, have confused the species with each other. Here, (a) misidentifications are corrected; (b) a neotype of Leucochrysa azevedoi is designated; (c) Leucochrysa (Nodita) morenoi, previously synonymized with Leucochrysa (Nodita) camposi, is recognized as a valid species [Reinstated status] All three species are redescribed and illustrated, with special emphasis on the types. Leucochrysa (Nodita) azevedoi was found to be relatively common in agricultural areas along Brazil’s Atlantic coast. The two other species are known only from their type localities: Leucochrysa (Nodita) camposi – coastal Ecuador, and Leucochrysa (Nodita) morenoi – Quito, Ecuador.

A New Genus and Species of Green Lacewings from Brazil (Neuroptera: Chrysopidae: Leucochrysini)

Abstract The adult and third instar of a new genus and new species of leucochrysine green lacewings, Santocellus atlanticis gen. et sp. nov., are described from Brazil’s Atlantic Forest. The first instar of Santocellus riodoce (Tauber) comb. nov. (formerly Leucochrysa riodoce) also is described. These two species share a distinguishing suite of adult and larval features: wings with dark (sometimes pustulate) markings; male with an elongate, cylindrical mediuncus and stiff membrane below; female with round, doughnut-shaped spermatheca having a shallow invagination and a sail-shaped velum that extends anteriorly; larvae with discrete, longitudinal head markings, short mandibles, broad labial palpi, and short, stubby, ventrally projecting terminal palpomeres. With this new genus (Santocellus gen. nov.), the number of genera in the tribe Leucochrysini increases to seven; Santocellus gen. nov. includes three described species: S. atlanticis sp. nov. and S. riodoce comb. nov., both from Brazil, and Santocellus bullata (Tauber) comb. nov., from Peru. Abstract RESUMO O adulto e a larva de terceiro instar de novo gênero e nova espécie de crisopídeo leucocrisíneo da Mata Atlântica do Brasil, Santocellus atlanticis gen. et sp. nov., são descritos. O primeiro instar de Santocellus riodoce (Tauber) comb. nov. (anteriormente Leucochrysa riodoce) também é descrito. Estas duas espécies compartilham um conjunto distintivo de características tanto do adulto como da larva: asas com manchas escuras (algumas vezes pustulosas); macho com mediuncus cilíndrico, alongado, com membrana rígida ventralmente; fêmea com espermateca arredondada, em forma de rosca, com uma invaginação rasa e com o velum grande, em forma de vela, estendendo-se anteriormente; larvas com manchas cefálicas separadas, alongadas longitudinalmente mandíbulas curtas, palpos labiais largos com palpômeros terminais curtos, grossos, projetando-se ventralmente. Com este novo gênero (Santocellus gen. nov.), o número de gêneros da tribo Leucochrysini aumenta para sete; Santocellus gen. nov. inclui três espécies descritas: S. atlanticis sp. nov. e S. riodoce comb. nov., ambas do Brasil, e Santocellus bullata (Tauber) comb. nov., do Peru.

Larvae of Four Leucochrysa (Nodita) Species (Neuroptera: Chrysopidae: Leucochrysini) from Brazil's Atlantic Coast

ABSTRACT Several morphological features and markings differentiate the larvae (three instars) of four Leucochrysa (Nodita) species (Neuroptera: Chrysopidae: Leucochrysini) that commonly occur in agricultural areas along the Atlantic Coast of Brazil. Descriptions and keys to species are provided. Based on the characteristics of the four species studied here and four previously described species, we propose a suite of diagnostic characteristics for Leucochrysa (Nodita) larvae.