Calopterygidae is a family of damselflies, in the superfamily Calopterygoidea. Members of Calopterygidae are commonly known as broad-winged damselflies, demoiselles, or jewelwings. These rather large damselflies have wingspans of 50–80 mm (compared to about 44 mm in the common bluetail damselfly, Ischnura elegans), are often metallic-coloured, and can be differentiated from other damselflies by the broader connection between the wings and the body, as opposed to the abrupt narrowing seen in other damselfly families. The family contains some 150 species.

The Calopterygidae are found on every continent except Antarctica. They live along rivers and streams.

Characteristics The adults have metallic bodies; their wings are broader, with wider bases than other damselflies, and at rest hold their wings parallel to the body, slightly elevated. Some species have conspicuously colored wings; in males, the wings are usually blue, without pterostigmata, in females green or brown. Species are often quite variable in color and patterning, and they are sexually dimorphic. Color intensity may fade with age. The wings are heavily veined, having often 18 or more antenodal veins. The first segment of their antennae is longer than the combined length of the other segments. They have a jerky, skipping form of flight similar to the flight pattern of a butterfly (fluttering, rather than hovering stably like many other damselfly and dragonfly families). At least one species of Calopterygidae has shown morphological plasticity in wing length associated with the closeness of a forest to the river or stream where it lives, with a more distant forest correlating to greater wing length. Calopterygids perch horizontally on twigs near the water's edge.

Calopterygidae nymphs have lateral gills are longer than the median gills. The nymphs have a flattened, pentagonal-shaped head, a long first antennal segment and long legs. They are found among submerged aquatic plants, woody debris and the exposed roots of streamside plants. There is a single generation per year. The time spent in the larval stage is influenced by both biotic factors, such as fat reserves, and abiotic factors, such as temperature, so they have the highest chances of surviving and reproducing.

Behavior

The mating system of most species in this family is resource defense polygyny, where males are often territorial, guarding riverine habitat that is sought after by females for egg deposition. Some males are not territorial. Within a species there may be a territorial and nonterritorial morph, which may be different in coloration.

Some species display courtship behavior, especially displays of wing movement by the male. At least one genus (Hetaerina) displays lekking behavior.

During mating, the male first removes other males' sperm from the female's reproductive tract, then places his own sperm there. The intromittent organ of the male has spines that physically remove rival sperm and also stimulate the female's muscles to contract and expel the sperm. In many species, the male accompanies the female when she searches for a site to lay eggs; in some cases, he even remains attached to her. The guarding of females post-copulation is done so another male does not mate with the female before laying her eggs even though the male may be able to reproduce with other females and in the case of Hetaerina species, the male may lose his territory during the time spent guarding.

Like all Odonata species, the species in this family are carnivorous in both their larval and adult stages. Larvae tend to feed on smaller invertebrates, such as mayflies.

Taxonomic history Calopterygidae was established by Selys in 1850 for damselflies related to the genus Calopteryx. It is one of the oldest family-group names in Odonata and has remained widely recognised throughout subsequent classifications.

During the twentieth century some authors proposed separating certain lineages from Calopterygidae, most notably the American Hetaerininae as the family Hetaerinidae. However, morphological and molecular studies consistently recovered Calopterygidae as a monophyletic lineage.

Modern classifications therefore retain a broadly defined Calopterygidae, with Hetaerininae treated as the sister group to all remaining members of the family.

Classification

Subfamiles and tribes according to Dijkstra et al. (2014)

Subfamily Calopteryginae – the demoiselles: Tribe Caliphaeini Caliphaea Tribe Calopterygini Atrocalopteryx Calopteryx Matrona Matronoides Neurobasis Tribe Iridictyonini Iridictyon Tribe Mnaisini Archineura Echo Mnais Psolodesmus Tribe Noguchiphaeini Noguchiphaea Tribe Saphoini Phaon Sapho Umma Tribe Vestalini Vestalaria Vestalis

Subfamily Hetaerininae – the rubyspots and others: Bryoplathanon Hetaerina Mnesarete Ormenophlebia

Fossil record Fossils attributed to Calopterygidae are known from the Eocene onward, with records from Europe, Asia and North America. Most fossil occurrences are from Oligocene and Miocene deposits in Europe.

Fossil genera The following fossil genera are currently placed in Calopterygidae: †Guangxicalopteryx †Sinocalopteryx

Etymology The family name Calopterygidae is derived from the type genus Calopteryx, with the standard zoological family suffix -idae.

The genus name Calopteryx is presumably derived from the Greek καλός (kalos, "beautiful") and πτέρυξ (pteryx, "wing"), likely referring to the broad and often brightly coloured wings of these damselflies.

References Calopterygidae. Digital Key to Aquatic Insects of North Dakota. Valley City State University.

Calopterygidae. Dragonflies and Damselflies of Ecuador. Electronic Field Guide Project, University of Massachusetts, Boston.

“Broad-Winged Damselflies.” Calopterygidae Family - Broad-Winged Damselflies, https://unsm-ento.unl.edu/Odonata/calo.html.

Dijkstra, K. B. Field Guide to the Dragonflies of Britain and Europe. British Wildlife Publishing, 2006. . Pages 23, 65.

Taylor, Philip D., and Gray Merriam. “Wing Morphology of a Forest Damselfly Is Related to Landscape Structure.” Oikos, vol. 73, no. 1, May 1995, p. 43. https://doi.org/10.2307/3545723.

Calopterygidae. UNH Center for Freshwater Biology.

Córdoba-Aguilar, Alejandro, and Adolfo Cordero-Rivera. “Evolution and Ecology of Calopterygidae (Zygoptera: Odonata): Status of Knowledge and Research Perspectives.” Neotropical Entomology, vol. 34, no. 6, Dec. 2005, pp. 861–879. https://doi.org/10.1590/s1519-566x2005000600001.

Córdoba-Aguilar, A., et al. (2009). The lek mating system of Hetaerina damselflies (Insecta: Calopterygidae). Behaviour, 146, 189–207.

Alcock, John. “Post-Copulatory Mate Guarding by Males of the Demselfly Hetaerina Vulnerata Selys (Odonata: Calopterygidae).” Animal Behaviour, vol. 30, no. 1, Feb. 1982, pp. 99–107. https://doi.org/10.1016/S0003-3472(82)80242-X.