Anup Shah / Eastern Chimpanzee (Pan troglodytes schweinfurthii) fourteen year old sub-adult male, named Fundi, making submissive facial gesture, Gombe National Park, Tanzania

Martin Willis / Budgerigar (Melopsittacus undulatus) father with begging chick at nest cavity, Queensland, Australia

Willi Rolfes/ BIA / Alpine Marmot (Marmota marmota) parent feeding with begging young, Hohe Tauern National Park, Austria

Eric Baccega / Pack of Grey wolves (Canis lupus) greeting one another and expressing submissive behaviour,captive, Bavarian Forest National Park, Germany, July. Captive.

Heike Odermatt / King Penguin (Aptenodytes patagonicus) chick begging for food, Saunders Island, Falkland Islands

Mark Taylor / Australian Shepherd pup, age 16 weeks, in play-bow stance.

Edwin Giesbers / Barbary macaque (Macaca sylvanus) baring teeth as a sign of submission, Gibraltar Nature Reserve, Gibraltar,

Suzi Eszterhas / Snowy Egret (Egretta thula) four week old chick begging for food, Sonoma County, California

Andy Rouse / Great crested grebe (Podiceps cristatus cristatus) courtship dance, Cardiff, UK, March.

Ole Jorgen Liodden / Waved albatross (Phoebastria irrorata) pair in courtship display, Espanola, Galapagos Islands. Critically endangered species.

Bruno de Faveri / Arctic Tern (Sterna paradisaea) with begging chick, Farne Islands, England, United Kingdom

Michael Quinton / Northern Flicker (Colaptes auratus) entering nest cavity with begging chicks, Alaska