Taxonomic Revisions of Six Genera of Entire-Eyed Owlflies (Ascalaphidae: Haplogleniinae), and First Large-Scale Phylogeny of the Owlflies



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The family Ascalaphidae, or owlflies, is a cosmopolitan and charismatic but poorly-understood and taxonomically ill-organized family of Neuroptera (lacewings). Prior to this work, it comprised approximately 450 valid described species placed in three subfamilies, 15 tribes, and 100 genera. In this dissertation, six genera of Haplogleniinae, or entire-eyed owlflies, are taxonomically revised based on cladistic analyses of morphological characters, and the first comprehensive phylogenetic analysis for the owlflies based on morphological and molecular characters is presented.

The taxonomic revisions are presented in three chapters, with their results as follows: (i) Allocormodes McLachan, a genus of cryptically-patterned species broadly distributed across tropical sub-Saharan Africa, is revised based on analysis of 35 morphological characters. Eleven species are recognized (five new), and one species name is synonymized. (ii) Tmesibasis McLachan, a genus distributed across sub-Saharan Africa and the southwestern Arabian Peninsula, is revised based on analysis of 39 morphological characters. Ten species (two new) are recognized as valid, and two species names are placed as synonyms. (iii) The entire-eyed owlflies of the Western Hemisphere are revised based on analysis of 79 morphological characters. Before this study, this group comprised 25 extant and two fossil species in six genera. Based on the cladistic analysis, a new classification is proposed, which places 37 extant species (13 new) in four genera: Amoea Lef?bvre, Ascalobyas Penny, Haploglenius Burmeister, and Neascalobyas new genus. Within Haploglenius, five new species groups are proposed, three of which represent former genera (Ascaloptynx Banks, Neohaploglenius Penny, and Verticillecerus van der Weele). Five species previously recognized as junior synonyms are also re-erected, and seven names are placed in synonymy with other species. In addition, Episperches molinai Nav?s and the fossil species Amoea electrodominicana Engel and Grimaldi are removed from Amoea and are placed within Ameropterus Esben-Petersen and Haplogleniinae incertae sedis, respectively. Ascaloptynx oligocenica Nel is also removed from the novel appendiculatus species group (formerly Ascaloptynx) within Haploglenius, and is placed within Haplogleniinae incertae sedis. For each of the genera treated, all species determined here to be valid are figured, keys to their identification are given, and maps of their distributions are provided.

In the fourth major chapter, the first large-scale phylogeny of the owlflies is presented, based on analysis of combined morphological (25 characters) and molecular (16S, 18S, and COI genes) datasets. These datasets were analyzed under maximum likelihood, Bayesian, and parsimony analytical regimes for 76 exemplars of Myrmeleontiformia (Ascalaphidae, Myrmeleontidae, Nemopteridae, Nymphidae, Psychopsidae), including 57 of Ascalaphidae. At the superfamily level, the families were recovered in all analyses in the form Psychopsidae + (Nymphidae + (Nemopteridae + (Myrmeleontidae + Ascalaphidae). Ascalaphidae was recovered as monophyletic in the Bayesian and parsimony analyses, and paraphyletic with respect to Ululodini and Myrmeleontidae in the maximum likelihood analysis. The subfamilies Haplogleniinae and Ascalaphinae were not recovered as monophyletic in any analysis. The Ululodini were monophyletic and well-supported in all analyses, as were the New World Haplogleniinae and the African/Malagasy Haplogleniinae. The remaining Ascalaphidae, collectively, were also consistently monophyletic, and include a genus traditionally placed in Haplogleniinae, Protidricerus van der Weele. None of the included tribes of non-ululodine Ascalaphinae were monophyletic in any analysis. Protidricerus was discovered to express a well-developed pleurostoma, a feature previously only encountered in divided-eye owlflies, and this feature may be important in future classifications. The feature traditionally used to differentiate the Haplogleniinae and Ascalaphinae, the entire or divided eye, can no longer be regarded as a reliable spot-diagnosis character to separate monophyletic groups within the family, and should be re-evaluated.